Flora (200 I) 196, 81 -100 http://www.urbanfischer.de/journalslflora © by Urban & Fischer VerLag Seed morphology in relation to infrageneric classification of Consolida (DC.) GRAY (Ranunculaceae) THEOPHANIS CONSTANTINIDIS*, GEORGE K. PSARAS, and GEORGIA KAMARI Section of Plant Biology, Department of Biology, University of Patras, GR-265 00 Patras, Hellas (Greece), E-mail: constgr@otenet.gr * Corresponding author Accepted: April 1, 2000 Summary The surface morphology of intact seeds of 37 taxa (species and subspecies) belonging to 6 different sections of the genus Consolida (Ranunculaceae) was investigated using scanning electron and light microscopy. The material studied covers the entire distribution range of the genus and represents c. 70% of the known Consolida taxa. Unlike certain species of the related genera Delphinium and Aconitum, all Consolida species have subpyramidal to subcylindrical seeds with pronounced transverse subparallel lamellae and testa cells covered with minute papillae. The crateriform hilum cavity is surrounded by well-formed or rudimentary lamellae, scales of various sizes or fringed appendages. The seeds of sect. Involutae, considered by some to represent the independent genus Aconitella, are subpyramidal and covered with low lamellae, which are usually expanded at the seed edges and proximal parts. Similar patterns are found in other Consolida species and splitting off the genus Aconitella is not supported, at least on the basis of seed micromorphology. In sect. Brevipedunculatae seeds are mostly subpyramidal and covered with scales of various sizes. Two informal groups are distinguished: the long-spurred allies of C. aucheri present a relatively shallow hilum furnished with dense, fringed appendages, while the hilum zone of the second, short-spurred group comprising the relatives of C. raveyi is surrounded by entire lamellae. The members of sect. Consolida also appear heterogeneous. Two subspecies of C. regalis (subsp. divaricata and subsp. paniculata) and C. tenuissima are unique on account of their constantly long and almost isolated scales. In contrast, the seeds of sect. Macrocarpae are uniform and do not permit distinctions among taxa. In sect. Longibracteolatae the seeds differ from other sections in having a subcylindrical shape and regular rows of complete, transverse, membranous rings. Based on similarities in seed morphology, it is proposed that the somewhat isolated C. lineolata should be included in sect. Longibracteolatae. After transferring C. barbata to sect. Involutae, sect. Parviflorae remains with a single species with exceptional floral morphology. It is concluded that seed morphology can be used in conjunction with inflorescence architecture, flower morphology and follicle characters as primary tool for infraspecific classification of Consolida. Seed morphology alone gives good support to three of the six sections of the genus. Key words: Consolida, Ranunculaceae, seed morphology, scanning electron microscopy (SEM), infrageneric classification. Introduction The genus Consolida (DC.) S. F. GRAY belongs to Ranunculaceae tribe Delphinieae SCHROD., together with Aco'nitum L. and Delphinium L. These three genera possess zygomorphic flowers, at least one spurred petal, and free follicles or a single follicle (TAMURA 1993). In its wide sense (i.e. including the members of the genus Aconitella SPACH - see below), Consolida comprises c. 52 species, less than the other two genera of the same tribe, each of which reaches or exceeds 300 species. 0367 -2530/01/196/02-081 $ 15.00/0 The taxonomic affinities of Consolida should be searched in Delphinium (TAMURA 1966), more precisely in the annual members of sections Delphinium and Anthriscifolium W. T. WANG (TRIFONOVA 1990). Taxonomic proximity and issues of nomenclatural compliance forced some authors to unite the two genera in the past (e.g. NEVSKII 1937). Besides morphology, close taxonomic relationship between Consolida and Delphinium is further revealed by karyological data (most members share the same basic number of n = 8, chromosomes ofR type; GREGORY 1941; TRIFONOVA 1973; BLANCHE et al. 1987), chemical evidence (presence of FLORA (2001) 196 81 particular patterns of seed fatty acids, AITZETMULLER et al. 1999; biosynthesis of distinctive diterpene alkaloids, also known to exist in Aconitum and Thalictrum, HEGNAUER 1990; HARBORNE & BAXTER 1993), and studies on chloroplast and ribosomal DNA (JOHANSSON 1995; Ro et al. 1997). Contrary to the vast majority of Aconitum and Delphinium species, Consolida always has an annual life cycle. Other unique Consolida features are the single spurred petal, probably resulting from merging of the two upper petals of Delphinium (SCHRODINGER, cited by TAMURA 1966), and the single follicle compared to 3 or 5 sessile follicles of Delphinium and 3 to several follicles of Aconitum. Consolida had early been recognized as a natural unit, at least since DE CANDOLLE' s time (1824). According to this author, it forms a section of Delphinium and comprises 11 species. BOISSIER (1867) subdivided Delphinium sect. Consolida based mainly on characters of the petal and spur. Altogether he recognized six Consolida groups marked with §, and increased the number of species to 32. A few years later HUTH (1895) monographed Delphinium and considered Consolida as a subgenus, further divided into six invalid "tribes". The taxa accepted reached 45; several of them cited as varieties today form widely recognized species. It was in the early 1920s that Soo (1922) transferred all European species of Delphinium with one petal and a single follicle to the genus Consolida, followed by MUNZ (1967 a, b), who treated the African and Asiatic species of the genus in the same way. The peculiar formation of the petal, upper sepal and spur of one subgroup of Consolida prompted KEMULARIA-NATHADSE (1939) to erect the new genus Aconitopsis, and to indicate similarities of her new genus with Delphinium, Consolida and Aconitum. More precisely, the eight species of Aconitopsis accepted are connected by the helmet appearance of the posterior sepal, gibbous, five-lobed petal and circinate spur ending. The name Aconitopsis was later rejected by SOJAC (1969), being replaced by Aconitella for reasons of nomenclatural priority. Simultaneously, the members comprising the genus rose to 10. Despite the accurate information provided for Consolida in both revision articles (Soo 1922; MUNZ 1967 a, b) and local floras (DAVIS 1965; IRANSHAHR 1992; CHATER 1993), the infrageneric classification of ~he genus is not yet clear and the taxonomic independence of Aconitella remains somewhat controversial. Seed coat micromorphology is known to be of considerable value for the taxonomy of Ranunculaceae subfamily Helleboroideae Hutch. (SEITZ 1969; MALYUTIN 1973; CAPPELETTI & POLDINI 1984; KARCZ & TOMCZOK 1987; BLANCHE 1990; MOLERO & PUIG 1990; ILARSLAN et al. 1997). Characters of the seed, inflorescence structure and flower morphology were used by 82 FLORA (2001) 196 MALYUTIN (1973, 1987) to evaluate phylogenetic schemes in Delphinium and propose a new infrageneric classification. Despite the importance of seed characters as a source of taxonomic information, no detailed studies on seed morphology of Consolida are known, with the exception of Trifonova (1984 a, b). The present study describes seed morphology in thirty-seven taxa of Consolida and simultaneously gives an overview of the variation encountered within the genus. Moreover, the information obtained by SEM and light microscopy is correlated with the classical infrageneric system of HUTH (1895), to evaluate its taxonomic efficiency. Materials and methods Plant material used in this study was collected either in the wild during excursions to Greece and Turkey or from herbarium specimens obtained on loan from E, LI, W, UPA (abbreviations according to HOLMGREN et al. 1990) and M. NYDEGGER-HuGLI. Particular importance was paid to countries such as Turkey and Iran where a large number of species exists. In contrast, extensive areas of central and western Europe comprise only one or two species, that may have been naturalized in post-mediaeval times (JALAS & SUOMINEN 1989). Precise collection data and localities are presented in the Appendix. Seed dimensions and crude information on seed morphology were obtained with the use of a Zeiss SV 6 stereoscope with a micrometric climax eye-piece. For each species at least 10 seeds were investigated, with the exception of C. aconiti where a lower number of seeds was available. Only crushed or immature seeds were found in herbarium specimens of C. saccata but when examined, their morphological proximity to other members of sect. Involutae was confirmed. For scanning electron microscopy, dry ripe seeds were mounted directly on stubs using double-sided adhesive tape and then sputter-coated with gold. Morphological observations were made with a Jeol6300 SEM. Images were recorded in digital form or as video printouts. To describe the seed coat sculpture, the terminology provided by STEARN (1992) and HARRIS & HARRIS (1994) is primarily used. Seed descriptions appearing in the major work on Antirrhineae by SUTTON (1988) were also particularly helpful. The infrageneric taxonomic scheme of HUTH (1895) is followed for the aggregation of the taxa in sections (Table 1), taken into consideration the annotations by TRIFONOVA (1990). Recently described taxa not dealt with by HUTH (1895) ':Yere placed together with their nearest relatives, with the exception of C. lineolata whose closest allies seemed obscure. Results Seeds of all the 37 Consolida taxa investigated have a subglobose or subpyramidal shape furnished with distinct lamellae that may vary in size, or with trans- Table 1. Sectional arrangement of Consolida taxa (species and subspecies) used in this study according to RUTH (1895). Number of investigated taxa in each section is given in parentheses. Underlined species have been relatively recently described and are not included in RUTH'S monograph. "Tribe" Propria has been replaced by sect. Consolida. Sect. Involutae (8 taxa) c. aconiti C. C. C. C. C. C. C. anthoroidea hohenackeri saccata scleroclada teheranica thirkeana stenocarpa Sect. Macrocarpae (4 taxa) C. C. C. C. ajacis brevicornis orientalis phrygia subsp. thessalonica Sect. Longibracteolatae (3 taxa) C. arena ria C. hellespontica C. olopetala Sect. Brevipedunculatae (10 taxa) C. C. C. C. C. C. C. C. C. C. aucheri axilliflora camptocarpa kabuliana kandaharica leptocarpa persica raveyi rugulosa tuntasiana Sect. Parviflorae (2 taxa) c. barbata C.flava Uncertain section (1 taxon) c. lineolata Sect. Consolida (9 taxa) c. glandulosa C. mauritanica C. oliveriana C. pubescens C. regalis subsp. regalis C. regalis subsp. divaricata C. regalis subsp. paniculata C. stallliana C. tenuissima verse and subparallel scaly rings. Seed dimensions, ornamentation, and scales or rings per seed side are summarized in Table 2. The hilum is situated at the lower part of the seed, opposite the apex of the pyramid, or at the wide part of the subcy lindrical seed. It is almost always hidden in a crateriform cavity and surrounded by scales. Seed surface observations for each section are described below. 1. Sect. Involutae [corresponding to HUTH's (1895) tribus Involuta, nom. illeg.] Eight species of this section were examined, i.e. Consolida anthoroidea (Fig. 1a -c), C. teheranica (Fig. Id-f), C. hohenackeri (Fig. Ig-i), C. thirkeana (Fig. lj-l), C. aconiti (Fig.2a-c), C. stenocarpa (Fig. 2d-f), C. scleroclada (Fig. 2g-i) and C. saccata. Of these, micrographs of C. saccata are not presented due to poor quality of the available seeds. However, their study revealed that they have the same morphological pattern as the rest of the section. Seeds more or less subpyramidal, with one convex side that comes in contact with the ± concave, inner part of the follicle surface (e.g. Fig. 1 g). Size between 1.5 x 1.1 and 2.7 xLI-I. 7 mm, colour light brown to dark brown, occasionally with different colour shades. Lamellae 11-21 per side, wavy, usually low and not overlapping (Fig. 1d, 2 g), or narrow and undulate (Fig. 2a, d), with a thickened (Fig. Ib) or blunt apex (Fig. I e), dirty white to yellowish, glossy and somewhat translucent (the greyish appearance of seed in some species is due to the reflecting effect of the paler lamellae). On edges and at the proximal and distal ends lamellae always much longer, c. 2-5 times as long as those of lateral faces (Fig. 1a, 2a). Almost continuous scale rings are found on the lower seed part, close to the crateriform hilum zone (Fig. lOa). Periclinal cell walls mostly convex (Fig. 1c) and finely granulate in all species examined (Fig. 1c, f, 2 i, 1); anticlinal walls straight and depressed (Fig. 2c, f). Small differences were observed among the eight species of sect. Involutae investigated. Consolida stenoFLORA (2001) 196 83 Table 2. Seed morphological characters in Consolida: sbpyr = subpyramidal, adaxial face convex, sbpyr/tetr = subpyramidal to tetrahedral, sharp edges, sbpyr/rd = subpyramidal, rounded edges, sbpyr/eon = subpyramidal to conical, sbgl = subglobose, sc!ed = scales that elongate at seed edge, erg = ± continuous scaly rings, se/und = undulate scales, sse = short scales not much projected from seed surface, rse = rows of disrupted scales, ise = ± isolate scales. Taxon Shape Dimensions (mm) C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. C. sbpyr sbpyr sbpyr sbpyr sbpyr sbpyr sbpyr sbpyr sbpyr/tetr sbpyr sbpyr sbpyr sbpyr sbpyr sbpyr/rd sbpyr sbpyr/rd sbpyr/tetr sbpyr/tetr sbpyr/rd sbpyr/rd subpyr/con subpyr/con subpyr/con sbpyr/tetr subpyr/con sbpyr sbpyr sbpyr sbpyr sbpyr/con sbpyrlcon sbpyr/con sbpyr sbpyrlcon sbgl 1.7 x 1.1 - 2.2 2.2 x 1.1 - 2.7 1.6 x 1.2 - 2.0 2.1 x l.3 - 2.7 1.9 x 1.1 - 2.3 1.8 x 1.1 - 2.2 1.S x 1.1 - 1.8 1.1 x 1.0 - 1.4 l.3 x 1.0 - 1.7 1.1 x 1.0 - 1.6 1.3 x 0.9 - 1.6 1.0 x 0.9 - 1.3 1.2 x 1.0 - 1.6 l.3 x 0.9 - 1.6 1.4 x 1.1 - 1.7 1.4 x 1.1 - 1.7 l.3 x 1.1 - 1.7 I.SxO.9-2.1 1.4 x l.1 - 2.1 1.4 x 1.2 - 2.0 1.8 x 0.9 -2.4 I.S x 1.0 - 2.1 1.4 x 1. 0 - 2.1 1.4 x 1.0 - 2.1 1.7 x 0.9 - 2.1 1.4 x 1.1 - 2.0 I.S x 1.1-2.1 1.4 x 1. 1 - 2.2 1.S x 1.1 - 2.2 1.S x 1.1 - 2.1 0.9 x 0.6- 1.2 1.1 x 0.9 - 1.7 1.7 x 1.1 - 2.0 2.0 x 1.1 - 2.7 0.9 x 0.8 - 1.1 0.9 x 0.9 - 1.2 aconiti anthoroidea hohenackeri scleroclada teheranica thirkeana stenocarpa aucheri axill ifl 0 ra camptocarpa kabuliana kandaharica leptocarpa persica raveyi rugulosa tuntasiana glandulosa mauritanica oliveriana pubescens regalis subsp. regalis regalis subsp. divaricata regalis subsp. paniculata stapfiana tenuissima ajacis brevicornis orientalis phrygia subsp. thessalonica arenaria hellespontica olopetala barbata C.flava C. lineolata carpa has the smallest seeds of the group (Fig. 2a-c). The seeds of C. thirkeana have an irregular, subpyramidal shape with rounded edges (Fig. lj). C. hohenackeri and C. thirkeana may occasionally have well-developed lamellae on their lateral sides. In this section however, development of lamellae is limited by the arrangement of seed in the follicle. Abaxial seed faces are in contact with adjacent seeds, while the adaxial side directly contacts the inner follicle wall. In both cases the development of long scales seems to be prevented. Welldeveloped scales appear on the transitional seed edges and proximal parts. On average, C. anthoroidea and C. scleroclada have the largest seeds of the group. The papillae that become evident at high magnification may 84 FLORA (2001) 196 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x l.3 1.7 1.S 1.7 1.S l.3 1.3 1.2 1.2 1.3 1.2 1.1 l.3 1.2 l.3 1.4 1.3 1.4 1.4 1.6 1.6 1.4 1.4 1.4 1.S 1.4 1.4 1.5 1.4 1.4 0.9 1.2 l.3 I.S 1.0 1.1 Ornamentation Number of scales/side sc/ed sc/ed sc/ed sc/ed sc/ed sc/ed sc/ed sc/ed crg sc/ed sc/ed sc/ed sc/ed sc/ed sc/und sc/ed sc/und rsc rsc sc/und crg rsc isc isc sc isc ssc ssc ssc ssc crg crg crg sc/ed rsc crg IS-20 IS-21 12-16 14-17 11-16 12-19 13-19 7-12 12-18 8-11 8-11 7-11 8-13 8-12 1O-IS 9-12 10-IS 11-14 8-12 10-14 II-IS II-IS II-IS II-IS 11-1S 11-16 12-16 12-16 II-IS l1-lS 8-12 9-13 9-12 16-20 6-9 8-11 cover the convex surface of longitudinal seed testa (c. aconiti, C. stenocarpa, Fig. 2c, f) or appear more locally on the grooves between cells (c. teheranica, Fig. If). 2. Sect. Brevipedunculatae (HUTH ex N. BUSCH) KEM.-NATH. Ten species of this section were examined, i.e. Consolida leptocarpa (Fig. 3 a -c), C. persic,'a (Fig. 3 d - f), C. kandaharica (Fig. 3 g - i), C. camptocarpa (Fig.3j-l), C. aucheri (Fig.4a-c), C. kabuliana (Fig.4d-f), C. rugulosa (Fig.4g-i), C. raveyi (Fig. 4j-I), C. tuntasiana (Fig. Sa-c) and C. axilliflora Fig. I. SEM micrographs of Consolida seeds: a, b, c, C. anthoroidea; d, e, f, C. teheranica; g, h, i, C. hohenackeri; j, k, 1, C. thirkeana. Magnification is noted on each micrograph. FLORA (2001) 196 85 Fig. 2. SEM micrographs of Consolida seeds: a, b, c, C. aconiti," d, e, f, C. stenocarpa," g, h, i, C. scleroclada," j, k, 1, C. barbatao Magnification is noted on each micrograph. 86 FLORA (2001) 196 (Fig.5d-f). With respect to flower morphology the section appears to be rather heterogeneous, with the short-spurred species of C. raveyi and its relatives forming one group, and the long-spurred allies of C. aucheri forming a separate group. Seeds more or less subpyramidal with a curved adaxial side (Fig. 3d, 4j), occasionally with somewhat rounded edges (c. tuntasiana, Fig. 5 a), size between 1.0 x 0.9 and 1.7 x 1.4 mm. Colour usually light brown, but may range from yellowish-brown to dark brown. Lamellae 7 to 18 per side (Fig. 3 g, 4 a), not forming distinct rings (with the exception of C. axilliflora, Fig. 5d) but discontinuous rows instead (Fig.4a, g), mostly of a straw-yellow colour, shiny, becoming elongate at edges (Fig. 3 j); endings smooth (Fig. 3 k, 5 e) or comb-like (Fig. 3 e, 4 e). Hilum zone in a crateriform formation, appearing variable among species. In certain species (c. axilliflora, C. raveyi, C. tuntasiana) surrounded by almost entire rows of lamellae, thus not differing from the pattern found in other Consolida sections (Fig. IOa, c), while in the rest of the Brevipedunculatae species (c. aucheri, C. camptocarpa, C. kandaharica, C. leptocarpa, C. rugulosa, C. persica) furnished with many narrow, fringe-like projections (Fig. lOb). Periclinal walls more or less convex (Fig. 3 f, 41), enriched with 2-3 /.tm granules of various density (Fig. 3 i, 4 i); anticlinal walls straight. An evident exception to the interrupted lamellae found in the majority of taxa of sect. Brevipedunculatae is C. axilliflora. In this species the lamellae form almost continuous rings (Fig.5d, e), which become slightly interrupted on the adaxial face and resemble patterns of sect. Longibracteolatae (see below). Other, less important deviations include the somewhat untidily placed lamellae of C. persica (Fig. 3 d) that do not form distinct rows, the characteristically long scales on edges and proximal parts of C. camptocarpa (Fig. 3j), and the subpyramidal seeds with rounded edges in the related species C. raveyi (Fig. 4 j) and C. tuntasiana (Fig. 5 a). C. kandaharica (Fig. 3 g) and C. camptocarpa (Fig. 3 j) have the lowest number of lamellae per seed side within the section. Further, the arrangement of papillae in C. axilliflora follows a specific pattern, leaving subparallel zones between lamellae apparently free of papillae (Fig. 5 e, f). 3. Sect. Consolida Seven species of this section were investigated with a total of nine taxa: Consolida pubescens (Fig. 5 g-i), C. mauritanica (Fig. 5j-l), C. glandulosa (Fig. 6a-c), C. oliveriana (Fig.6d-f), C. stapfiana (Fig.6g-i), C. regalis subsp. regalis (Fig. 7 a-c), C. regalis subsp. divaricata (Fig. 7 d - f), c. regalis subsp. paniculata (Fig. 7g-i) and C. tenuissima (Fig. 7j-l). Morphologi- cal characters of the seeds appear heterogeneous among species, and include taxa with almost complete rings of lamellae (c. pubescens, Fig. 5 g), taxa with pronounced, elongate scales (c. regalis subsp. paniculata Fig. 7 g ; C. tenuissima, Fig. 7 j) and intermediate forms. Seed shape subpyramidal with rounded edges (Fig. 6a, g) to subcylindrical (Fig. 5 j); in C. stapfiana and C. glandulosa with sharp and flat faces that give seeds a subtetrahedral appearance. Size between 1.4 x 0.9 and 2.4 x l.6 mm. Colour may vary from light grey-brown (c. oliveriana) to dark blackish-brown (c. glandulosa, C. regalis group). Lamellae 11 to 15 per side (with the exception of C. mauritanica), well separated from each other by sinuses of various depth (Fig. 5 k, 6 e), in precise transverse rows (Fig. 6 d, 7 a), not overlapping (Fig. 5 g, 7 b), becoming longer and more robust at edges and proximal parts, usually whitish-grey, straw yellow, dirty yellow or brownish. Lamellae occasionally crossed by clear longitudinal striae (Fig. 5 h, 7 e) that indicate separation lines between two successive testa cells. In some species (c. mauritanica, Fig. 5 k; C. oliveriana, Fig. 6 e) lamellae with a dilated and blunt ending and a somewhat sinuate margin. In C. pubescens, contrary to other taxa, the lamellae form almost complete rings (Fig. 5 g, h) interrupted locally by deep sinuses. Three taxa, i.e. C. regalis subsp. divaricata (Fig. 7 d, e), C. regalis subsp. paniculata (Fig. 7 g, h) and C. tenuissima (Fig. 7 j, k) are unique in having very large, discreet and elongate scales that overlap each other and give the impression of an echinate seed. Hilum zone situated in a crateriform cavity that is quite shallow in the C. regalis group and C. tenuissima. The same zone may also be quite wide (c. glandulosa) or may be indicated by a darker colour, compared to the rest of seed surface, as in C. oliveriana. Periclinal walls more or less convex, furnished with small papillae of various density (Fig. 6 f, 7 i); anticlinal walls straight, sometimes with sparse papillae (Fig. 71). 4. Sect. Macrocarpae (HUTH ex N. BUSCH) KEM.-NATH. Four taxa of section Macrocarpae were available for study, namely Consolida ajacis (Fig. 8a-c), C. orien- • talis (Fig.8d-f), C. phrygia subsp. thessalonica (Fig. 8g-i) and C. brevicornis (Fig. 8j-I). Seeds subpyramidal, often with one (rarely more) distinctly convex side (Fig. 8 a). Size between 1.4 x 1.1 and 2.2 x 1.5 mm. Colour dark brown to blackish. Transverse lamellae 11 to 16 per side, short, patent and wavy (Fig. 8 b, e, h), pale brown to yellowish, usually paler than main body, becoming more pronounced and less undulate on edges and at the hilum area. Hilum zone FLORA (2001) 196 87 Fig. 3. SEM micrographs of Consolida seeds: a, b, c, C. leptocarpa; d, e, f, C. persica; g, h, i, C. kandaharica; j, k, 1, C. camptocarpa. Magnification is noted on each micrograph. 88 FLORA (2001) 196 Fig. 4. SEM micrographs of Consolida seeds: a, b, c, C. aucheri,' d, e, f, C. kabuliana,' g, h, i, C. rugulosa,' j, k, I, C. raveyi. Magnification is noted on each micrograph. FLORA (2001) 196 89 Fig. 5. SEM micrographs of Consolida seeds: a, b, c, C. tuntasiana; d, e, f, C. axilliflora; g, h, i, C. pubescens; j, k, 1, C. mauritanica. Magnification is noted on each micrograph. 90 FLORA (2001) 196 Fig. 6. SEM micrographs of Consolida seeds: a, b, c, C. glandulosa; d, e, f, C. oliveriana; g, h, i, C. stapfiana; j, k, I, C. ./lava. Magnification is noted on each micrograph. FLORA (2001) 196 91 Fig. 7. SEM micrographs of Consolida seeds: a, b, c, C. regalis subsp. regalis .. d, e, f, C. regalis subsp. divaricata .. g, h, i, C. regalis subsp. paniculata .. j, k, 1, C. tenuissima. Magnification is noted on each micrograph. 92 FLORA (2001) 196 Fig. 8. SEM micrographs of Consolida seeds: a, b, c, C. ajacis; d, e, f, C. orientalis; g, h, i, C. phrygia subsp. thessalonica; j, k, I, C. hrevicornis. Magnification is noted on each micrograph. FLORA (2001) 196 93 Fig. 9. SEM micrographs of Consolida seeds: a, b, c, C. olopetala; d, e, f, C. hellespontica; g, h, i, C. arenaria; j, k, 1, C. lineolata. Magnification is noted on each micrograph. 94 FLORA (2001) 196 Fig. 10. SEM micrographs of the hilum area of C. teheranica (a), noted on each micrograph. situated in a crateriform depression, mostly surrounded by rudimentary lamellae (Fig. 8 g), occasionally with a slight projection at its centre. Periclinal testa walls convex, usually covered with papillae along their main axis (Fig. 8 c, 1); anticlinal walls straight. In some species zones that lacked papillae were observed (Fig. 8f, 1). The seeds of all species in sect. Macrocarpae look alike and distinction among taxa using seed characters is difficult. Variation of seed ornamentation is also low; the same seed pattern was observed in all species and all populations examined. 5. Sect. Longibracteolatae HUTH ex ThIF. Three species belonging to this section were studied, i.e. Cansolida olopetala (Fig. 9a-c), C. hellespontica (Fig. 9d-f) and C. arenaria (Fig. 9g-i). Seeds conical, rarely subpyramidal (Fig. 9 a, d, g); one side sometimes convex. Size between 0.9 x 0.6 and 2.0 x 1.3 mm, colour dark brown to blackish. Lamellae usually well-developed (less so in C. arenaria), in entire, transverse rows (Fig. 9b, e), not differentiated into isolated scales, sometimes completely covering seed surface (Fig. 9b), dirty yellow to brownish and shiny. Hilum area located in a crateriform depression (Fig. lOc), surrounded by entire and curved rows of lamellae, with minute, fringe-like projections. In C. hellespantica the central part of the hilum area may form a characteristic apiculate appendage. This formation, although extant, is less apparent in the other two species of this section. Periclinal walls more or less convex and finely granulate (Fig. 9c, f); anticlinal walls straight and slightly raised. In the case of C. arenaria papillae of two different sizes: these on testa ridges sparse and almost double the size of those on the elongated and shallow furrows between ridges (Fig. 9h, i). c. leptocarpa (b) and C. hellespontica (c). Magnification is 6. Sect. Parviflorae HUTH ex TRIF. The two species of this section seem to be taxonomically unrelated and their seeds deserve separate descriptions. Seeds of Consolida barbata (Fig.2j-l) large (2.0 x 1.1 to 2.7 x 1.5 mm), subpyramidal, with one distinctly convex size (Fig. 2j), brownish. Lamellae in transverse, interrupted rows, usually shorter on the faces and more elongate on the edges, proximal, and occasionally distal part, dirty yellow to pale brownish and shiny; endings somewhat blunt and dilated (Fig. 2k). Hilum area hidden in a crateriform cavity and surrounded by curved, almost entire lamellae. Periclinal walls indicated by longitudinal striae, convex and covered with clear papillae (Fig. 21). Seeds of C. flava (Fig.6j-l) small (0.9 x 0.8 to 1.1 x 1.0 mm), mostly broadly subconical, light brown. Lamellae disrupted, forming distinct subparallel rows, yellowish, shiny and translucent (Fig.6j). Hilum zone in a crateriform cavity, surrounded by fringe-like projections as in species of sect. Brevipedunculatae. Small papillae clearly observed on testa cells (Fig. 61). 7. Uncertain section The original description of C. lineolata (HUBERMORATH 1978) and sequential annotations (DAVIS et al. 1988) do not indicate any close relative in the genus. Accordingly, this species is examined separately (Fig.9j-I). Seeds small (0.9 x 0.9 to 1.2 x 1.1 mm), subglobose and brownish, completely covered by long, imbricate lamellae (Fig. 9j). Lamellae in complete rows, sometimes disrupted by shallow sinuses, or by the slightly unequal length of the testa cells (Fig. 9k), pale yellowish and shiny, edges somewhat fringed. Hilum zone in a FLORA (2001) 196 95 basal depression, surrounded by curved lamellae, central part usually slightly projected. Periclinal walls more or less convex and finely granulate (Fig. 91); anticlinal walls straight and somewhat depressed. Discussion Compared to its allies, Consolida is an evolved genus with precise synapomorphies (reduction of carpels from three or more to one, complete loss of lateral petals, spur consisting of one petal) that are not found in any other species of Delphinium and Aconitum. Most Consolida species are adapted to the Mediterranean type climate or more arid climate types of the Irano-Turanian zone (ZIMAN & KEENER 1989). Pronounced periods of drought in these areas have certainly favoured the exclusive annual life cycle of Consolida. The biogeography of the genus indicates that Turkey, in particular Anatolia (c. 29 taxa) should be considered as the centre of diversity, with further radiation of species into the Irano-Turanian area (c. 23 taxa), Greece (c. 10 taxa) and countries around the Mediterranean. Consolida forms a coherent, monophyletic clade with Delphinium and Aconitum. Some authors propose a direct evolution line of Consolida from Delphinium (TAMURA 1966, MALYUTIN 1973). This phylogenetic evidence combined with its advanced morphological characters and present distribution area, make it reasonable to assume that most of Consolida's evolutionary process has taken place in the Eastern Mediterranean/ Orient from a common Aconitum/Delphinium ancestor. In Delphinium, three major seed structures have been reported (MALYUTIN 1987). The c. 170 perennial species of D. subgen. Delphinastrum (DC.) PETERM. are characterized by seeds with marginal wings (type 1, semina alata vel subalata), which are not covered by scales in transverse rows of any kind, although elongated processes of single testa cells or small groups of cells have very occasionally been reported [D. hansenii (E. GREENE) E. GREENE, WARNOCK 1990]. The small group of D. subgen. Staphisagria (DC.) PETERM. lacks wings, lamellae or scales on its seeds, which exhibit a reticulate-rugulose surface pattern (type 2, semina reticulato-rugulosa, see also ILARSLAN et al. 1997). In contrast, the numerous perennial members of D. subgen. OZigophyllon DIMITROVA (c. 175 species) and the c. 15 species of the annual subgen. Delphinium have seeds characterized by distinct rows of lamellae (type 3, semina squamosa). None of the 37 Consolida taxa examined in this study displays seed characters of type 1 or 2. As seed morphology has been used as a primary tool to delimit presumably natural groups in Delphinium, the unique seed type 3 of Consolida may serve as supporting evidence that connects this genus with parti96 FLORA (2001) 196 cular branches of Delphinium. This assumption may be applicable in cases where seed characters have been evolved once, and needs to be strengthened by additional phylogenetic evidence. In tracing Consolida's ancestors we agree with TRIFONOVA (1990) in considering its evolution from the annual members of D. sect. Anthriscifolium as highly improbable. Section Anthriscifolium is geographically confined to the eastern Asiatic region (BLANCHE 1990), far beyond the Anatolian centre of diversity for Consolida. The easternmost known Consolida species, C. schlagintweitii (HUTH) MUNZ, is recorded from the area of Kashmir (MUNZ 1967 b); while no species of Consolida occurs in East Asia. Seed morphology of the related genus Aconitum seems to be more complex. The unispecific subgen. Gymnaconitum (STAPF) RAPCS. from Central Asia comprises the only annual species of Aconitum, which bears lamellate seeds (TAMURA 1966). In subgen. Lycoctonum (DC.) PETERM. all sections are characterized by lamellate seeds except of the monotypic sect. Alatospermum TAMURA that has alate seeds (LAUENER & TAMURA 1978, TAMURA & LAUENER 1979). In the taxonomically problematic subgen. Aconitum, no precise seed character-state can be combined with other morphological characters to delimit particular groups. However, observations by SEM on A. anthora L. from Europe have related the alignment of round-top tubercles occurring on adjacent seed testa cells with the development of distinct transversal ridges that may progressively expand into membranous wings (MOLERO & PUIG 1990). This insight could shed light on the ontogeny of lamellae through a procedure not necessarily confined to Aconitum but also characterizing Delphinium and Consolida. Some recent phylogenetic work based on evidence derived from epidermal micromorphology (HOOT 1991) and molecular data (HOOT 1995, JENSEN et al. 1995), has associated Nigella L. with the zygomorphic Ranunculaceae group which includes Consolida. Available information on seed coat structure of six Nigella species (KARCZ & TOMCZOK 1987) however, revealed several morphological dissimilarities with Consolida, especially in primary seed ornamentation and testa cell shape. Within Consolida, the most prominent group comprises the species of section Involutae. All species of this section have the same seed pattern of low discontinuous ridges that expand into well-formed scales mostly along seed edges and proximal parts. The testa is covered with granules of different density that become clearly visible at a magnification of x 200 or higher. The comparatively low variation in seed morphology within sect. Involutae indicates a stable sectional unit that has even been erected into generic rank (Aconitella, see TRIFONOVA 1990), albeit without gaining general acceptance. Apart from common seed morphology, the members of section Involutae present similarities in flower morphology and inflorescence structure, and share a more advanced petiolar anatomy (TRIFONOVA 1990). However, the delimitation of the group as a whole appears somewhat ill-defined. Consolida barbata, traditionally placed in sect. Parviflorae, exhibits the typical pentamerous partition of spurred petal as in the members of sect. Involutae, but neither a hood formation of petal nor a circinate petal ending. Seed coat morphology favours the placement of C. barbata in sect. Involutae, a practice already proposed by KEMULARIA-NATHADSE (1939); SOJAC (1969) and TRIFONOVA (1990). By transferring C. barbata to sect. Involutae, the section itself becomes more inhomogeneous, leaving only a pentamerous spurred petal to be used as a discriminating floral parameter. When the variable petal formation in Consolida (petal in C. olopetala lacking lateral lobes ; petal of C. flava with low lateral lobes and a tetrahedral4-merous or 5-merous central lobe, see under sect. Parviflorae) and the drastic changes in floral structure among Aconitum, Delphinium and Consolida s. 1. are taken into consideration, then generic rank of sect. Involutae as Aconitella seems unjustified. Seed morphology of this section serves to connect its members through a common seed pattern but similar patterns are also found in other Consolida species. Nevertheless, floral and inflorescence morphology of sect. Involutae appear to represent a distinct evolutionary trait within Consolida, and it is therefore recommended that sub generic rank be maintained for this section. In Consolida sect. Brevipedunculatae the seeds are mostly subpyramidal with rounded edges. They are furnished with rows of well-defined transverse lamellae that do not form distinct rings, with the exception of C. axilliflora. In this group two different lines can be distinguished, based mostly on flower morphology and further supported by seed microcharacters. The first line covers several species centred in the area of Iran and Afghanistan and radiating out into the neighbouring areas of Pakistan, Turkmenistan, Central Asia and Iraq, reaching Syria and Turkey in the west. The members of this line (seven of which were examined, i.e. C. aucheri, C. camptocarpa, C. kabuliana, C. kandaharica, C. leptocarpa, C. persica, C. rugulosa) are characterized by flowers with long, often ascending spurs and a superficial overall resemblance to the annual members of Delphinium sect. Delphinium. Their seed coat morphology is particularly homogenous. The fringed appearance of scales surrounding the hilum area (Fig. 1Ob) serves as a distinguishing character that was observed in at least nine populations of the six species examined, with the exception of C. kabuliana, where more seeds need to be investigated, preferably from more than one population. The second line of sect. Brevipedunculatae covers species distributed further west, from Syria and Turkey to Greece. They have flowers with very short pedicels and short spurs not exceeding petal length. Of the three species examined (c. axilliflora, C. raveyi, C. tuntasiana), C. axilliflora differs in having a subtetrahedral seed shape and almost continuous transverse rings on seed faces, whereas seeds of the other two species are round with interrupted scales. This difference in seed morphology is difficult to evaluate. It would be of interest to examine C. cruciata, an ally of C. axilliflora, to determine its seed sculpture. The members of this line do not present the striking fringed appearance of the hilum zone, although minute projections resembling fringes can be found around the hilum. The taxa belonging to C. sect. Consolida exhibit a series of seed ornamentation that ranges from the relatively undifferentiated, entire rings of the western Mediterranean C. pubescens, to the almost isolated long scales of the Greek endemic C. tenuissima. Most of the species examined connect these two extreme forms by having short subparallel lamellae with deep sinuses and occasionally with undulating margins. The long, disconnected scales of three taxa (c. regalis subsp. divaricata, C. regalis subsp. paniculata and C. tenuissima) are unique in the genus; however, the weakly divided lamellae of C. regalis subsp. regalis arranged in rows connect these extreme forms with other members of the section. Two series have been proposed within sect. Consolida, based mainly on pedicels characters, flower morphology and karyological evidence (BLANCHE et al. 1987). The two members of series Pubescentes (c. mauritanica and C. pubescens) have rather dissimilar seed sculpture but share three metacentric chromosome pairs in their complement, in opposite to the examined members of series Consolida that have only two metacentric chromosome pairs. Seed morphology of all four taxa of C. sect. Macrocarpae indicates a profoundly coherent group with dark, subpyramidal seeds bearing low, wavy lamellae in transverse, subparallel series. Rudimentary lamellae are found around the hilum zone. The uniformity of the group is further supported by flower morphology, inflorescence architecture, chromosome studies and ecological preferences. This section has the widest distribution area of all sections, its taxa spreading from the western Mediterranean to central Asia. Expansion of distribution is certainly favoured by human activities, as all the species are weeds of cultivated land. Section Longibracteolatae is characterized by remarkable uniformity in its seed coat characters. Unlike other Consolida sections, all the species of sect. Longibracteolatae have subcylindrical seeds with continuous lamellae forming partly overlapping rings, with the exception of C. arena ria, which has rather short rings. FLORA (2001) 196 97 This particular seed morphology is limited to sect. Longibracteolatae and approaches seed structure of several species of the annual Delphinium sect. Delphinium. Members of sect. Longibracteolatae all possess hairy follicles put on pedicels refracted downwards (evident on at least the lower part of the inflorescence), and a particular chromosome morphology characterized by the lack of a second metacentric or submetacentric chromosome pair which is common in many other Consolida species (CONSTANTINIDIS & KAMARI, unpublished). The same type of seed morphology was observed in C. lineolata, a recently described and little known species from Turkey. Apart from similarities in seed morphology, C. lineolata shares the same hairy follicles growing on deflexed pedicels as other members of sect. Longibracteolatae. This evidence allows safe placement of C. lineolata in sect. Longibracteolatae, where it is obviously the most deviating representative. After transferring C. barbata to sect. Involutae, the only member of sect. Parviflorae examined is C.flava. Its seeds, apart from their small size, show no obvious difference from those of other members of C. sect. Consolida or sect. Brevipedunculatae, however its flower morphology is quite distinct. This species has very short lateral lobes on its spurred petal, while the middle part appears tetrahedral and divided into four or five parts by shallow notches. The corolla segments are yellow tinged with purple spots in their inner parts. This flower structure appears quite unique in the genus and permits, at least temporarily, the inclusion of C. flava in its own section. To conclude, it is found that seed morphological characters of Consolida give support to three of the six infrageneric groups recognized by HUTH (1895), i.e. sects Involutae (better erected to subgeneric rank), Macrocarpae and Longibracteolatae. Sections Brevipedunculatae and Consolida are more diverse in both seed and flower features and may need further segregation. A preliminary evaluation of taxonomic criteria in Consolida based on gross morphology reveals that inflorescence architecture, floral structure and follicle characters can provide additional evidence for an infrageneric classification of the genus. As a strict annual, Consolida possesses no way of population maintenance other than seed distribution and successful germination. The various species of the genus present different breeding systems (DE-YUAN J 986; BOSCH et al. 2001; CONSTANTINIDIS unpublished) and a noteworthy specialization for ecological preferences. Although surface sculpture of plant organs has been ascribed particular ecological/evolutionary roles (BARTHLOTT 1981), the biological significance of seed surface in Consolida has been largely ignored. 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Appendix Material examined for seed morphology_ C. aeoniti (L.) LINDLEY - Turkey, C3 Afyon, Strasse Dinar Denizli, 10 km nach Dinar, zwischen Dinar und Dazkiri, felsige kahle Steppe, 980 m, 21. 07.1975, Simon 75 109 (BASBG). c. ajaeis (L.) SCHUR - a) Greece, Nomos EtoliasAkarnanias, c. 7 km SW Navpaktos, field margins, 60 m, 21. 06.1998, Constantinidis 8543 (UPA). b) Greece, Nomos Argolidos, W of Didima village, abandoned fields, 200 m, 06.06.1998, Constantinidis 8547 (UPA). c. anthoroidea (BOIss.) SCHROD. - a) Turkey, Provo Van, Van - Geva~, FLORA (2001) 196 99 c. 15 km from Van, stony hillside, 2000 m, 02.09.1956, McNeill 724 (E). b) Iran, Provo Luristan, Dorud, openings among oakbushes, 5000 ft., 23. 08. 1941, Koelz 18498 (W). c. arenaria CARLSTROM - Greece, Nomos Dodekanisou, Rodos island, S ofCharaki settlement, sandy seashore, 0-5 m, 06.07.1998, Constantinidis 8577 (UPA). C. axillijlora (DC.) SCHROD. - Turkey, C7 Adiyman, W Kar~dg, Brache, 900 m, 30.05.1983, Sorger 83-18-2 (W). C. aucheri (Bmss.) IRANSHAHR - Afghanistan, Provo Herat, in valle fluvii Heri Rud infra Chisht (Tschischt), ca. 1500 m, 04.08.1962, Rechinger 19206 (W). C. barbata (BUNGE) SCHROD. - a) Afghanistan, Kala Sarkari, 7000 ft., 09. 09. 1939, Koelz 13935 (W). b) Afghanistan, Provo Takhar, Khost-o-Fereng, unteres Fereng-Tal, Chahar Qarya, 1700 m, 20. 07. 1965, Podlech 11948 (E). C. brevicornis (Vis.) Soo - Greece, Nomos Kefallinias, Kefallinia island, close to Agios Gerasimos Omalon, fields, 400 m, 16. 06. 1998, Constantinidis 8545 (UPA). C. camptocarpa (FISCH. & C.A. MEY. ex LEDEB.) NEVSKI - a) Iran, Tehran - Mashhad road, low hills along the road, 50 miles east of Shahrud, 08. 07. 1972, Alava 10876 & Iranshahr (E). b) Afghanistan, bei Maimana, Neubauer 4242 (E, W). c) Afghanistan, Obey springs, 100 km east of Herat, on dry slopes, 2100 m, 13.07.1969, Andersen & Petersen 406 (E). C. flava (DC.) SCHROD. - a) Iraq, Haswa desert, 20 km west of Rutba, Ramadi Liwa, 08. 06. 1957, Rechinger s. n. (E). b) Iraq, distr. Baghdad (Mesopotamia), Jazira, in deserto arenoso 36 km a Tuz Kurmatli versus Indiyana, 22. 06. 1957, Rechinger 10612 (W). C. glandulosa (Bmss. & HUET) BORNM. - a) Turkey, B5 Kayseri, 3 km S Incesu, Salzsteppe, 1000 m, 16.07.1969, Sorger 69-60-10 (W). b) Turkey, Cappadocia, close to Akkoy village, roadsides, 1280-1350 m, 31.07.1999, Ylldmmh & Constantinidis (Herb. Ylldmmh). C. heUespontica (Bmss.) CHATER - a) Greece, Nomos Fthiotidos, c. 0.5 km SE of Grammeni village, scree and rocky slopes along road side, ophiolithic substrate, c. 250 m., 8. 06. 1997, Constantinidis 7069 (UPA). b) Greece, Nomos Trikalon, c. 2.8 km after the village of Kalomira along road to Mourgani, serpentine scree along road side, c. 580 m, 10.08.1998, Constantinidis 8000 (UPA). C. hohenackeri (Bmss.) GROSSH. - Turkey, A9 Kars, 10 km S Sarikamis, offener Pinus sylv. Wald N der Strasse, 2200 m, 23.07.1981, Sorger 81-48-5 (W). c. kabuliana (AKHTAR) IRANSHAHR - Afghanistan, Kabul, Gulbahar, Amsel s.n. (W). C. kandaharica IRANSHAHR Afghanistan, Kandahar, in lapidosis 10-20 km NE Kandahar, 1100 m, 26.05.1967, Rechinger 35241 (W). c. leptocarpa NEVSKI - a) Afghanistan, Provo Baghlan, mittleres AndarabTal, zwischen Kushdarrah und Mushi, LoBhange, 11. 06. 1965, Podlech 11307 (E). b) Afghanistan, Provo Qataghan, lower Andarab valley, to 14 km E. of village, roadside, 1100 m, ol. 07. 1965, Lamond 2299 (E). c) Turkmenistan, peripheria oppidi Ashabad, loco Keshi dicto, 350-500 m, 14.09. 1976, Vasak s. n. (W). c. lineolata HUB.-MoR. & SIMON - Turkey, C4 I~el, Ermenek - Mut, 43 km SW Mut, Pinusaufforstung auf Kalk, 940 m, 22.07.1978, Nydegger 13313 (E). C. mauritanica (COSSON) MUNZ - Spain, Guadalajara, Cillas, en un barbecho camino hacia Torrubia, 1160 m, 2l. 07. 1996, Pisco & Martinez s.n. (MA). C. oliveriana (DC.) SCHROD. - Iraq, Distr. Mosul (Kurdistan), ad confines Turkiae provo Hakari, inter Mosul et Zakho, in collibus siccis, 02. 07. 1957, Rechinger 100 FLORA (2001) 196 10627 (W). C. olopetala (Bmss.) HAYEK - Turkey, Provo Erzincan, Erzincan - Kelkit, c. 12 km from Erzincan, dry hillsides, 1500 m, 01. 08.1957, Davis & Hedge 31909 (E). C. orientalis (GAY) SCHROD. - a) Turkey, A6 Sivas, Sariyar (NO Circir), Feldrand, Bachrand, 1500-1700 m, 15.07. 1969, Sorger 69-57-27 (W). b) Iran, Provo Shahrud-Bustam, in declivibus australibus montium Shahvar ad Nekarman (Nigarman), 1000 m, 20-26.07.1948, Rechinger 6248 (W). C. persica (Bmss.) SCHROD. - a) Iran, Provo Hamadan, 137 km from Hamadan on the road to Ghazvin, sandy and stony mountain slope, 2170 m, 18.07.1974, Assadi & Amini 13626 (W). b) Iran, Fars, Shiraz region, hillside (and edges of cultivated fields) along the road to Bushir, 32 km from Shiraz, 26.06. 1972, Alava 10642 (E). c) Iran, Tehran, Evin, 07. 1965, Mirzayan 6604E (W). C. phrygia (Bmss.) Soo subsp. thessalonica (Soo) P. H. DAVIS - Greece, Nomos Trikalon, ESE Vasiliki village, cultivated fields, 130 m, 28. 06. 1998, Constantinidis 8544 (UPA). C. pubescens (DC.) Soo - Spain, Caroli Pau Herbarium Hispanicum, Peiiegoloza, 26. 07. 1934 (MA). C. raveyi (Bmss.) SCHROD. - a) Turkey, ProVo Ankara, above Tuz GOlii, 25 km north of Ko~hisar, stony slopes - rock ledges, c. 925 m, 29.07.1956, McNeill 336b (E). b) Turkey, Anatolia, W of Kmkale, margins of abandoned fields, 1050 m, 28.07.1999, Ylldmmh & Constantinidis (Herb. Ylldmmh). C. regalis subsp. divaricata (LEDEB.) MUNZ - Azerbaijan, 8 km S of Miyanduab, semi-desert, 1200 m, 09. 07.1964, Grant 16074 (W). C. regalis subsp. paniculata (HOST) Soo - a) Greece, Nomos Florinis, close to the village of Trigonon, cultivated fields and field margins, 950 m, 12.08.1998, Constantinidis & Garofalo 8541 (UPA). b) Greece, Nomos Etolias Akamanias, c. 2.3 km S Frangouleika village, Klisoura gorge, among bushes, 130 m, 06.08.1998, Constantinidis K241 (UPA). c. regalis S. F. GRAY subsp. regalis - Croatia, InnerIstrien, NW Lupoglav, N Buzet, an der StraBe nach Martin, Ackerrain, 80 m, 27.07.1997, Starrniihler s. n. (W). C. rugulosa (Bmss.) SCHROD. - Iran, Provo Kashan, Mashhad Ardahal, 24.05.1970, Iranshahar 13541-E (W). C. saccata (HUTH) DAVIS - Iraq, Dohuk, Mosul Liwa, beside path among vine fields, 02.10.1957 (?), Haines 1218 (E). C. scleroclada (Bmss.) SCHROD. var. rigida (FREYN & SINT.) DAVIS - Turkey, Provo Tunceli, Tunceli, 1100 m, 23. 07.1957, DAVIS & HEDGE 31592 (E). C. stapfzana DAVIS & SORGER - Turkey, C3, 20 km SW Korkuteli, Feldrand, 1200 m, 12.07.1968, Sorger 68-2711 (LI). c. stenocarpa (DAVIS & HOSSAIN) DAVIS - Turkey, C3 Antalya, 20 km SW Korkuteli, Steppenreste, 1200 m, 12.07.1968, Sorger 68-27-9 (W). c. teheranica (Bmss.) RECH. fil. - a) Iran, Provo Mazanderan, in valle fluvii Talar infra Abbasabad, ca. 1200 m, 04.07.1937, Rechinger 2015 (W). b) Iran, Mazanderan, Haraz valley above Panjab, 1400 m, Ol. 08.1959, Wendelbo 1651 (E). c) Iran, ProVo Elbourz, Gadouk, Komrand to Roudbar, 24. 07. 1948, Behboudi. & Aellen 5378E (W). C. tenuissima (SM.) Soo - Greece, Nomos Attikis, Mt. Pateras, slopes of Kandili low summit, limestone, 350-500 m, Ol. 05. 1994, Constantinidis 4499 (UPA). C. thirkeana (Bmss.) SCHROD. - Turkey, Dikmen s. Ankara, 07.09.1954, Auner s. n. (W). c. tuntasiana (HAL.\cSY) SooGreece, Nomos Attikis, Mt. Gerania, Makriplagi summit, limestone, 1340-1350 m, 07.05.1995, Constantinidis & Iliadis 5507 (UPA).