identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03D57350FFD2227AD207BEA72F549E11.text	03D57350FFD2227AD207BEA72F549E11.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ablepharus Fitzinger 1824	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> 4.1. On  Ablepharus taxonomic issues, phylogeny and hemipenial morphology </p>
            <p> Akin to  Mabuya (S´anchez-Martínez et al., 2020),  Ablepharus suffers from nomenclatural and taxonomical issues rooted in taxonomic descriptions based on external morphological characters (colouration, size, pholidosis, etc.; see Mertens, 1952; Fuhn, 1969a,b, 1970; G¨oçmen et al., 1996; Schmidtler, 1997) and related to the small number of examined specimens (usually from only a few localities). The scarcity of diagnostic characters and the lack of sufficient data on morphological variation results in difficulties regarding the interpretation of some taxa. Nevertheless, based on the published literature Karamiani et al. (2021b) synthesised a few of the external morphological characters and created an identification key to the Iranian species of the genus. However, the key contains some generalizations, which cause practical difficulties in species determination using the respective characters (e.g., “prefrontals usually forming a median suture”). Based on our observations of numerous (over 400)  A. kitaibelii specimens (unpublished data), many of the traditionally used diagnostic characters either partially overlap or are highly variable even within a single population (e.g., number of supralabials, scales around the midbody, nuchals, etc.). As a result, the number of taxa within  Ablepharus is still disputed. For example, Uetz et al. (2023) and Karamiani et al. (2021b) treat  A. budaki anatolicus as a species –  A. anatolicus . This assumption is probably rooted in a few molecular studies (Poulakakis et al., 2005; Skourtanioti et al., 2016; Bozkurt and Olgun, 2020) which have revealed a basal taxon from the Island of Kastellorizo, Greece and the area around the Turkish Town of Kaş. It is likely that the specimens from the region have been assigned to this taxon by presumption. Understandably, none of these works provides a nomenclatural act of raising the subspecies to species. The authors of these publications consider the data inconclusive and claim that additional specimens are needed. Nevertheless, the "  A. anatolicus " combination continues to persist in both the published literature and in internet sources. By assigning  A. bivittatus to the genus  Asymblepharus, Bozkurt and Olgun (2020) provided a new combination and generated even further confusion. Additional complications stem from the assignment of some species to different subfamilies. These issues demonstrate the need for a thorough revision not only of  Ablepharus but all allied lineages as well (Shea, 2021). The boundaries of the genus  Ablepharus were recently expanded by the synonymisation with  Himalblepharus and  Asymblepharus (Mirza et al., 2022) . The current taxon  Ablepharus remains poorly defined from a morphological point of view as it was never formally re-described and a diagnosis was never formulated. The present definition of  Ablepharus is based mainly on molecular phylogeny (Mirza et al., 2022). That said, it should be emphasised that the ablepharine skinks with completely fused eyelids form a well-supported monophyletic clade nested within  Ablepharus (Mirza et al., 2022) . Further taxonomic studies may recover its subgeneric or even generic status. </p>
            <p> The phylogenetic position of the fused eyelids representatives of the genus  Ablepharus was much more controversial in the past, particularly considering the pioneering study of Pyron et al. (2013), in which  A. budaki and  A. kitaibelii plus  A. chernovi formed a well-supported clade with a set of lygosomine species, while  A. pannonicus was grouped with  Asymblepharus alaicus Elpatjevsky, 1901 into a relatively distant clade sister to a large “  Sphenomorphus group” radiation. This phylogeny clearly makes the genus  Ablepharus paraphyletic. In a similar manner, Datta-Roy et al. (2013) recovered the clade  Ablepharus grayanus plus  Asymblepharus himalayanus (Günther, 1864) as sister to the “  Sphenomorphus group”. Macey et al. (2006) demonstrated the close relationship between  A. pannonicus and the members of the genus  Asymblepharus [represented by  A. alaicus and  A. sikimmensis (Blyth, 1854) ], which formed a well-supported clade. In addition to molecular data, the representatives of the genus  Asymblepharus were grouped by a set of morphological traits (Jeriomtshenko, 1987; 2002), although the existence of this genus was neglected in some studies (see Shea and Greer, 2002) and the species were commonly attributed to the genus  Scincella . Based on external morphology, Jeriomtshenko and Szczerbak (1986) regarded  A. grayanus and  A. pannonicus as closely related. The phylogeny of Mirza et al. (2022) (generally repeated by Liang et al., 2024), resolved many issues with the “ablepharine” genera and recovered the monophyly of the clade. According to their phylogeny,  Ablepharus alaicus and  Ablepharus eremchenkoi (Panfilov, 1999) (both formerly in  Asymblepharus ) are basal to the fused eyelids ablepharine skinks. Sister to their clade is the clade of the former (now assigned to  Ablepharus )  Asymblepharus species :  A. nepalensis ,  A. mahabharatus ,  A. sikimmensis ,  A. cf. ladacensis ,  A. himalayanus . However, the available phylogenies of the group are based on various genetic markers from different taxa, e.g., ND 2, 12S, 16S, etc. (Shea, 2021) and on very different taxon sampling (some taxa are represented by a single sample), and therefore cannot be accepted unequivocally. According to Jeriomtshenko and Szczerbak (1980),  A. lindbergi is morphologically related to  Asymblepharus . No molecular data is available for  A. lindbergi and  A. darvazi . To date, there have been no molecular studies based on type material. </p>
            <p>The animal genitalia are rapidly diverging structures (Eberhard, 1985). Traditionally, in certain animal groups, including reptiles, the morphology of genitalia has been used for creation of taxonomic and sometimes phylogenetic hypotheses (e.g., Keogh, 1999; Zaher, 1999; Nunes et al., 2014). An alignment of a cluster based on hemipenial morphology with a phylogeny based on other characters (e.g., molecular) is an intriguing approach, and may reveal discrepancy between divergence of a certain structure and divergence of the species.</p>
            <p> The grouping (Fig. 4A), based on slender and robust lobes, does not overlap in general to the molecular phylogeny of Mirza et al. (2022) (Fig. 4B); phylogenetically,  A. pannonicus is closer to  A. deserti and  A. kitaibelii is a sister taxon to  A. chernovi (contrary to the morphological cluster). The relationship between  A. budaki and  A. rueppellii , is supported by both molecular phylogeny and morphology of the hemipenes. </p>
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	https://treatment.plazi.org/id/03D57350FFD2227AD207BEA72F549E11	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Vergilov, Vladislav;Zlatkov, Boyan	Vergilov, Vladislav, Zlatkov, Boyan (2024): Morphology of hemipenes and its taxonomic implication in the fused eyelids species of the genus Ablepharus (Squamata: Scincidae). Zoologischer Anzeiger 312: 79-91, DOI: 10.1016/j.jcz.2024.07.009, URL: https://doi.org/10.1016/j.jcz.2024.07.009
03D57350FFD12276D151BEC82CFD9F0C.text	03D57350FFD12276D151BEC82CFD9F0C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scincidae	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> 4.3. On  Scincidae phylogeny and hemipenial morphology </p>
            <p>At present, hemipenial morphology data representing all main phylogenetic clades (sensu Pyron et al., 2013) is available for approximately 3 % of all scincid species. These data help us outline a preliminary picture of the main trends in hemipenial evolution in this poorly studied lizard family.</p>
            <p>Many studies, dealing with hemipenes, simply describe the morphology of these structures without providing figures or often present the structures only schematically. This is likely the main reason for generating a plethora of terms for homologous structures. Sometimes a complete eversion of the hemipenis is not easily achieved, which has resulted in an additional source of confusion in some works (e.g., Zhang, 1986).</p>
            <p> Greer (1979) recognised two basic types of hemipenes (illustrated schematically on figs. 11–15) in Australian skinks, and this discrimination can be accepted for the whole family: 1) with relatively short columnar base and slightly bulbous or bilobed cap; and 2) with long and narrow base and two equally long bifurcations. The “short columnar base” is typical for the  Egernia group which includes the genera  Egernia ,  Tiliqua and  Tribolonotus . The genus  Cyclodomorphus and  Corucia zebrata Gray, 1855 can be assigned to the same type (Ziegler and Bohme ¨, 2004). The close relationships between the members of this group are supported by molecular phylogenetic analyses (Gardner et al., 2008; Pyron et al., 2013). The “short columnar base” type also includes the  Eugongylus group (sensu Greer, 1979) and its two subgroups, i.e., the </p>
            <p> Eugongylus subgroup with the genera  Anotis, Cryptoablepharus ,  Emoia ,  Eugongylus ,  Leiolopisma, Morethis , and  Proablepharus , and the  Lampropholis subgroup with the genera  Carlia ,  Lampropholis , and  Menetia . S´anchez-Martínez et al. (2020) studied the hemipenes of a number of  Mabuya and  Trachylepis taxa. Seven of the  Mabuya species have thick hemipenes with a very short bifurcation:  M. agilis ,  M. altamazonica ,  M. caissara ,  M. bistriata ,  M. dorsivittata ,  M. frenata ,  M. macrorhyncha . Similar short bifurcation is observed in  Trachylepis atlantica (Schmidt, 1945) , and  Tiliqua rugosa (Gray 1825) . Noble and Bradley (1933) provide drawings of the hemipenes of  Plestiodon fasciatus L. which appear rather similar to those of  T. capensis (the present study). The hemipenes are characterised by wide lobes, numerous folds on the lobes, short bifurcation and base, and the presence of an asulcal protrusion (lateral aspect according to fig. 12 caption in Noble and Bradley, 1933) similar to that observed in  Ablepharus budaki (Fig. 3F).  Scincella baraensis Nguyen, Nguyen, Nguyen &amp; Murphy, 2020 has a very simple, balloon-shaped, non-bifurcated hemipenis without folds and protrusions (Nguyen et al., 2020).  Riopa guentheri (Peters, 1879) also possesses a non-bifurcated hemipenis, but it is possible that the hemipenes were not completely everted and that the tip is actually not visible (Fig. 3 in Bhilala et al., 2021). Bifurcation is lacking in  Copeoglossum nigropunctatum (Spix, 1825; S´anchez-Martínez et al., 2020). </p>
            <p> According to Greer (1979), the second type of hemipenes is deeply bifurcated with a long narrow base and two equally long lobes. This type is characteristic of the Sphenomoprhus group (after Skinner et al., 2013) including  Anomalopus ,  Ctenotus ,  Eremiascincus ,  Hemiergis ,  Lerista ,  Notoscincus ,  Saiphos ,  Sphenomorphus , and  Concinnia queenslandiae (De Vis, 1890) . This condition is considered derived, in contrast to the columnar structure condition in the genera  Eumeces and  Mabuya which is considered basal. The latter statement is not valid for  P. fasciatus (Noble and Bradley, 1933) and  Eutropis longicaudata (Ziegler, 2002) , which possess a weak hemipenial bifurcation (type one, according to Greer, 1979), or some species with a substantial degree of bifurcation (type two, op. cit.) such as  Eutropis multifasciata (Zhang, 1986) (species previously attributed to genus  Eumeces and  Mabuya ). In  Tropidophorus noggei Ziegler, Thanh &amp; Thanh, 2005 , the hemipenes are weakly bifurcated (Ziegler et al., 2005) although the genus  Tropidophorus is shown to be closely related to  Sphenomorphus species by Pyron et al. (2013). An extreme bifurcation of the hemipenial lobes is found in  Scincella rufocaudata (Darevsky &amp; Nguyen van Sang, 1983; Darevsky and Nguen, 1983) in which the lobes are approximately three times longer than the base. However, this condition may due to an incomplete eversion of the pedicel (the base). The general hemipenial shape, i.e., having fine lobes as long as the base, of  Scincella reevesii (Gray, 1839) (Ziegler, 2002) is similar to the hemipenes of  A. kitaibelii and  A. budaki described by Vergilov et al. (2017). In  Sphenomorphus buenloicus (Darevsky &amp; Nguyen van Sang, 1983) the lobes are shorter than the base, whereas in  S. indicus (Gray, 1853) they are longer than the base (Zhang, 1986; Ziegler, 2002). In  S. incognitus (Thompson, 1912) (Zhang, 1986) , the length of the lobes cannot be estimated due to incomplete eversion. The genus  Sphenomorphus is paraphyletic, with  Scincella deeply nested within it, and the hemipenial morphology of the groups supports this point of view (Shea, 2012; Linkem et al., 2010; Linkem, Diesmos and Brown, 2011). The species of  Ablepharus examined in this study can also be assigned to the deeply bifurcated hemipenes type. </p>
            <p>These examples demonstrate that the degree of bifurcation of the hemipenis, although well-defined, is an extremely homoplastic character.</p>
            <p> Scincid hemipenial morphology has received relatively little attention and has been studied rather sporadically which may explain the inconsistency in terminology. The folds of the organ caused tremendous confusion and the lack of understanding of their homology resulted in the coining of numerous terms. One of the first authors dealing with  Scincidae hemipenes was Cope (1896), who noticed the presence of "longitudinal laminae" in many scincids.  S. reevesii (Ziegler, 2002) has longitudinal folds on the sides of the base, not reaching the branches. No longitudinal folds were observed in the  Ablepharus species. Cope (1896) mentioned some "cross-ribbed plicae" in  E. carinata and  P. obsoletus . Noble and Bradley (1933) found a series of "transverse flounces" on the side opposite to the "heart-shaped pad" in two scincid lizards. These, however, may be artifacts of sample processing, i.e., injection of salt solution and fixation with formaldehyde. McCann (1949) also mentions “transverse plicae” on the apical surface of the lobes in  E. carinata . S´anchez-Martínez et al. (2020) observed “horizontal ridges” in  T. atlantica . These folds are apparently homologous to the transverse folds of  T. capensis (Fig. 1E). The term "horizontal" is inappropriate from a morphological point of view. Apparently, the anatomical position of the everted hemipenes is different from those of the isolated organs (oriented to the taste of the researcher). For this reason, the terms “transverse folds” or “transverse plicae” should be considered correct. </p>
            <p> Lateral folds (Fig. 3) can be seen on the lateral part of the hemipenial base in many  Scincidae species. The term “lateral” in this case is inappropriate, because of the variable orientation of the everted organ of a living animal. Dunger (1973) mentioned "flap-like lobules" on the hemipenis of  Leptosiaphos kilimensis (Stejneger, 1891) from Nigeria (now attributed to  Leptosiaphos dungeri Trape, 2012 ). The lateral projections observed in  Hemiergis gracilipes (Greer 1979) are likely homologous to the "bulbous lobes" of  A. budaki (Vergilov et al., 2017) , although they are substantially more pronounced in the latter. The term "bulbous lobes" is confusing, as the term "lobes" has been used for the branches of the bifurcated hemipenis by other authors. For us, due to the lack of more appropriate term for such structures, the usage of term “lateral folds” or “lateral plicae” is acceptable. Greer (1989) mentioned that the genera  Carlia ,  Lampropholis ,  Lygisaurus , and  Saproscincus share a unique elongated projection of the base. This character can be an autapomorphy of the group, which forms a well-supported clade in the phylogenetic tree of Pyron et al. (2013). Linkem et al. (2011) referred to the lateral region of the "main shaft" basad of the hemipenial bifurcation as "bulbous lobe", treated it as an autapomorphy of the clade, and described the new genus  Pinoyscincus . These “lobes” appear homologous to the lateral folds (as those of  A. kitaibelii ; Fig. 2) but are wider, thicker and cover the whole lateral surface of the hemipenial base. </p>
            <p> The sulci are bordered by thick folds for which we encourage the use of the term “labia” (Fig. 3) (these structures correspond to the “sulcal lips” of Klaver and Bohme ¨1986). The labia extend to the sulcal openings at the apices of the lobes where they transform into small elongated projections, i.e., the “terminal awns” (e.g., Montingelli et al., 2022). Vergilov et al. (2017) referred to these structures as the “terminal awls [sic]”. In  Trachylepis and  Mabuya , S´anchez-Martínez et al. (2020) referred to the labium as the “fold of the base of the sulcus spermaticus”. The figure labelling of this “fold” by S´anchez-Martínez et al. (2020) corresponds to the base of the labia of the hemipenes (where the structure is thicker) but does not include the apex of the lobes. Cope (1896) describes the labia (we assume that the author refers to this structure) as “welt” that is “opposite the sulcus spermaticus”. </p>
            <p> The term “asulcal protrusion” was proposed by Vergilov et al. (2017) for  A. budaki , it was observed in  A. rueppellii (this study), and in all cases it is homologous with a similar structure in  T. capensis . This structure has also been observed in other species. S´anchez-Martínez et al. (2020) referred to this structure as the “rounded fold”. Noble and Bradley (1933) presented a similar protuberance on the asulcal surface of the hemipenis of  Eumeces fasciatus , calling it a “tubercule”. A similar but less protruded fold (very similar to the one in  T. capensis ), is present in  Pinoyscincus abdictus abdictus (Brown &amp; Alcala 1980) (Linkem et al., 2011) . Presumably, all these structures are homologous and apparently represent a homoplastic trait. </p>
            <p> Thе use of inappropriate terminology is not limited to the hemipenial folds and has been applied to variety of other structures. For example, Vergilov et al. (2017) divided the hemipenis into apex, truncus, and pedicel based on the terminology of Klaver and Bohme ¨(1986) for  Chamaeleonidae . This terminology, however, appears incompatible with the scincid hemipenis, which differs substantially. </p>
            <p> There are a number of examples regarding the usefulness of hemipenial morphology in support of phylogenetically close relationships. Bohme ¨(1988) pointed out the similarity in hemipenial morphology in ecologically distinct genera, such as the specialised semiaquatic genus  Cophoscincopus and the morphologically unspecialised  Leptosiaphos ianthinoxantha (B¨ohme, 1975). Both taxa possess a collar-like ring, which was interpreted as a synapomorphy by the author. This finding likely indicates a close relationship between the two taxa and was partly supported by Trape et al. (2012). B¨ohme (1988) also mentioned the close similarity between  L. ianthinoxantha and the species present in Dunger (1973), here interpreted as  L. dungeri , both of which have bifurcated hemipenes. Bohme ¨et al. (2000) included figures of the hemipenes of the closely related  Cophoscincopus greeri B¨ohme, Schmitz, &amp; Ziegler, 2000 and  C. durus (Cope, 1862) . The authors mentioned that the organs were densely covered with delicate pustules except the whole apical area, however, they did not discuss the nature of these structures. </p>
            <p>No scincid species have been previously reported to possess calcified structures, as also shown in the present study. Their absence can, to some extent, be used in the general diagnosis of scincid lizards. Noble and Bradley (1933) noted that the hemipenes of scincids and iguanids are less complex than those of teiids and lacertids which are, in relation to mating behaviour, more generalised in the first two groups. Although this field remains largely unexplored, the driving force of sexual selection has already been partly discussed (Eberhard, 1985; 2010).</p>
            <p> The usage or interpretation of the specific hemipenial morphology for higher taxonomic analyses, however, is less informative, e.g., the lizard family  Lacertidae (B¨ohme, 1971). This is mainly due to numerous homoplastic traits found in non-congeners (for example the asulcal protrusion in  A. budaki and  T. capensis ; Fig. 1E; 2B, C). Similarly, homoplastic characters can be pointed out for relatively closely related boid snakes (Ziegler and Bohme ¨, 1997). This leads to the conclusion that hemipenial differentiation should be used only after a stable phylogeny of a group of interest has been established. In this respect, satisfactory results for the family  Gymnophtalmidae were presented by Nunes et al. (2014). </p>
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	https://treatment.plazi.org/id/03D57350FFD12276D151BEC82CFD9F0C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Vergilov, Vladislav;Zlatkov, Boyan	Vergilov, Vladislav, Zlatkov, Boyan (2024): Morphology of hemipenes and its taxonomic implication in the fused eyelids species of the genus Ablepharus (Squamata: Scincidae). Zoologischer Anzeiger 312: 79-91, DOI: 10.1016/j.jcz.2024.07.009, URL: https://doi.org/10.1016/j.jcz.2024.07.009
