An updated list of the Mexican herpetofauna: with a summary of historical and contemporary studies.

The growth in our knowledge of the diversity of the herpetofauna of Mexico has occurred over the period of approximately 445 years from the work of Francisco Hernández to that of a broad multinational array of present-day herpetologists. The work of this huge group of people has established Mexico as one of the most significant centers of herpetofaunal biodiversity in the world. This status is the result of a complex orography, in addition to diverse habitats and environments and the biogeographic history of Mexico. The current herpetofauna consists of 1,421 native and introduced species, allocated to 220 genera, and 61 families. This figure is comprised of 1,405 native species and 16 non-native species (as of April 2023). The non-native species include two anurans, 13 squamates, and one turtle. The level of endemism is very high, presently lying at 63%, with this level expected to increase with time. Species richness varies among the 32 federal entities in the country, from a low of 50 in Tlaxcala to a high of 492 in Oaxaca. Amphibian species richness by state-level can be envisioned as comprising three levels of low, medium, and high, with the lowest levels occurring in the Peninsula of Baja California, a group of seven states in north-central and central Mexico, and a group of three states in the Yucatan Peninsula, with the highest levels occupying the southern states of Guerrero, Puebla, Veracruz, Oaxaca, and Chiapas, and the medium level in the remaining states of the country. Reptile species richness also can be allocated to three categories, with the lowest level occupying Baja California Sur, a group of central states, and the states of the Yucatan Peninsula, and the highest level found in a cluster of the states of Veracruz, Guerrero, Oaxaca, and Chiapas. Knowledge of the Mexican herpetofauna will continue to grow with additional studies on systematics, conservation, and the construction of checklists at various levels.

Does size matter? An analysis of the niche width and vulnerability to climate change of fourteen species of the genus Crotalus from North America.

The niche comprises the set of abiotic and biotic environmental conditions in which a species can live. Consequently, those species that present broader niches are expected to be more tolerant to changes in climatic variations than those species that present reduced niches. In this study, we estimate the amplitude of the climatic niche of fourteen species of rattlesnakes of the genus Crotalus to evaluate whether those species that present broader niches are less susceptible to the loss of climatically suitable zones due to the projected climate change for the time period 2021-2040. Our results suggest that for the species under study, the breadth of the niche is not a factor that determines their vulnerability to climatic variations. However, 71.4% of the species will experience increasingly inadequate habitat conditions, mainly due to the increase in temperature and the contribution that this variable has in the creation of climatically suitable zones for most of these species.

Influence of microhabitat use on morphology traits of three species of the Anolis sericeus complex (Squamata: Dactyloidae) in Mexico.

The relationship between microhabitat use and morphology in Anolis lizards has been well studied in the Caribbean islands. However, studies of ecomorphology are scarce for the vast majority of mainland Anolis species. Thus there is a great lack of knowledge on how microhabitat use may influence the morphology of most mainland Anolis species. In this study we evaluated the relationship between morphology and microhabitat use in three sister species of the genus Anolis (A. sericeus, A. unilobatus and A. ustus) inhabiting the mainland and examined whether sympatric coexistence with other Anolis species affects microhabitat use and morphology of Anolis sericeus. The results of this study showed that A. ustus perches on higher, thinner, and warmer branches than its sister species (A. sericeus and A. unilobatus), and that snout-vent length, arm, forearm and femur length and weight of the three species are positively correlated with perch diameter, while the number of subdigital lamellae and weight are slightly correlated with perch height. In addition, the absence of sympatric Anolis species increases the variability in perch height, weight, arm and forearm length, and hip width observed in A. sericeus. Differences in limbs length, head length, weight and number of lamellae enabled the identification of two groups of species exploiting statistically different microhabitats. The relationship between microhabitat use and morphology indicates that some morphological traits, such as locomotor structures, weight and number of subdigital lamellae of the three species are influenced by microhabitat use and that interspecific competition may reduce variability in the height of perches used.

NATURAL HISTORY OF THE BLACK IGUANA Ctenosaura similis (SQUAMATA: IGUANIDAE) IN ISLA CONTOY, QUINTANA ROO, MEXICO / Historia natural de la iguana negra Ctenosaura similis (Squamata: Iguanidae) en isla Contoy, Quintana Roo, México

ABSTRACT The genera Iguana and Ctenosaura belong to the Iguanidae family, and populations of most species of these genera have decreased due to anthropogenic effects. The natural history of most species of this family is poorly known, including Ctenosaura similis. Therefore, this study documents some ecological aspects of the species, such as its feeding habits, and habitat and microhabitat use in a population of Isla Contoy in Quintana Roo, Mexico. The data showed that even though C. similis is distributed throughout the island, individuals more commonly use human construction (buildings) of Parque Nacional Isla Contoy (PNIC) and mangrove areas. Microhabitats most used by all age classes and sexes were rocks and trees. The diet of the species is omnivorous; however, the frequency of prey type consumed changes with the age of the individuals (hatchlings, juveniles, and adults), as well as with the season of the year. The practice of cannibalism by some individuals is also reported, as well as coprophagy and scavenging. This work provides general information on the life history of C. similis, complementing the scarce information that is currently available.

RESUMEN Las iguanas de los géneros Iguana y Ctenosaura son algunas de las especies de escamados que han tenido una importante disminución en sus poblaciones naturales, principalmente por factores antropogénicos. No obstante, a pesar de esto existen especies de las que el conocimiento sobre su historia natural es escaso, como es el caso de la iguana negra Ctenosaura similis. Por lo que, en este trabajo se documentan algunos aspectos de la ecología, tal como los hábitos alimentarios y uso del hábitat y del microhábitat de la población de la iguana negra (C. similis) en Isla Contoy, en el estado de Quintana Roo, México. Los análisis de este estudio revelaron que a pesar de que C. similis se distribuye en toda la isla, utiliza con mayor frecuencia las instalaciones del Parque Nacional Isla Contoy (PNIC) y la zona de manglar; los microhábitats más usados por las diferentes clases de edad y los diferentes sexos, fueron las rocas y los árboles. En general, la dieta de esta especie es omnívora, pero la frecuencia con la que consumen ciertas categorías de presa cambia con la edad del individuo (cría, juvenil y adulto), así como a través de las estaciones del año. Además, se observó que algunos individuos practicaron el canibalismo, la coprofagia y la carroñeria. Este trabajo aporta datos generales de la historia natural de C. similis, lo que complementa la escasa información que hasta ahora se tiene disponible.

An updated checklist of the herpetofauna of Querétaro, Mexico: species richness, diversity, and conservation status.

Mexico contains a large number of amphibian and reptile species, and the states in the southeastern part of the country are the richest and most diverse. Although the study of species richness within Mexico's individual states has been increasing over the last two decades, herpetofaunal species lists for several states in the central region are incomplete. Herein, we provide a list of the herpetofauna of the state of Querétaro, a state that remains relatively unexplored. We also indicate the conservation status of the component species in the state, based on the national and international categorizations, as well as their Environmental Vulnerability Scores (EVS). Based on a review of literature and specimens in scientific collections, our results show that the herpetofauna of Querétaro currently is composed of 138 species, of which 34 are amphibians (27 anurans and seven caudates) and 104 are reptiles (three turtles, 34 lizards, and 67 snakes). The number of Mexican endemic species occurring in the state consists of 19 amphibians and 50 reptiles, representing 55.9% and 48.1%, respectively, of the total numbers in these groups. A total of 61 species (13 species of amphibians and 48 of reptiles) are listed in NOM-059-SEMARNAT-2010, within the categories of Special Protection (Pr) and Threatened (A). According to the International Union for Conservation of Nature (IUCN 2019), 34 species of amphibians (100% of species) and 87 species of reptiles (83.65% of a total of 104) have been placed within a risk category, i.e., except for the DD and NE categories. The EVS results show that nine species of amphibians have high environmental vulnerability, 11 species medium vulnerability, and 14 low vulnerability, whereas among the reptiles, 22 species have high vulnerability, 43 medium vulnerability, and 37 species low vulnerability. The herpetofauna of Querétaro illustrates a high level of species richness, which is typical of central Mexico, as well as diverse groupings associated with the arid, temperate, and tropical environments found in the state. These types of studies are necessary for increasing our knowledge of the herpetofauna in the various states of Mexico, as well as for evaluating the conservation status and implementing management plans for species in both herpetofaunal groups.

Comparative morphology and trophic ecology in a population of the polymorphic lizard Sceloporus minor (Squamata: Phrynosomatidae) from central Mexico.

Polymorphism among individuals of the same population has generally been linked to alternative reproductive tactics, where different morphs can exhibit differences in their morphological, ecological, and behavioral attributes. These differences may result in a divergence in diet between morphs due to differential exploitation of habitat, morphological differences that influence prey selection, or differential energy expenditure that results in different nutritional needs. The present study analyzes the morphology (morphometry and body mass) and diet of red and yellow male morphs in a population (El Enzuelado) of the lizard Sceloporus minor from central Mexico. No differences between morphs were found for any of the morphometric variables analyzed (snout-vent length, tail length, jaw length, jaw width, head length, head width, head height, tibia length, femur length, forearm length and ventral patch length). In both morphs, allometric growth was observed in all body features analyzed, as well as in morphometric features of the head across seasons. Analysis of stomach contents showed that the diet of red males was composed of 12 categories of prey, while that of yellow males was composed of 10 categories; those categories of diet not shared between morphs (e.g., Isoptera, Psocoptera) were consumed by their respective morph in very low proportions. Categories of diet with the highest values of food importance for both groups were Coleoptera, Orthoptera, and leaves; a similar pattern was seen across seasons. This, in turn, is reflected in low niche breadth values for each morph and a very high niche overlap. There were no significant differences between morphs overall, or between morphs per season, in the weight and volume of stomach contents or in the number of prey items found in stomachs; however, differences in these variables across all males (independent of morph) were recorded between seasons. Likewise, no significant correlations were found between body size (snout-vent length) and the volume of stomach contents for either morph or between lizard mandibular dimensions and the volume of stomach contents for red morph males. For the yellow morph, prey volume unexpectedly decreased significantly with jaw size rather than increasing as expected. Overall, this study adds new information about the morphology and feeding of males in this species, and suggests that in this population, color morphs lack the morphological and ecological differences found in some other species of polymorphic lizard.

Ecology of the growth of Anolis nebulosus (Squamata: Dactyloidae) in a seasonal tropical environment in the Chamela region, Jalisco, Mexico.

Juvenile growth rates are thought to be restricted by available food resources. In animals that grow throughout the year, such as tropical lizards, growth is therefore predicted to be faster during the rainy season. We test this prediction using a population of Anolis nebulosusby describing the growth trajectories of both sexes using nonlinear regression models, and we then correlate the growth rates of individuals with food available in the environment, precipitation, and temperature. The Von Bertalanffy model fits the growth rates of the females better, while the logistic-by-length model fits the males better. According to both models, the males grew faster than females, reaching slightly smaller sizes at adulthood. Males reached sexual maturity when 35 mm long, at an age of seven months, and females matured at 37 mm (SVL), taking nine months to reach this size. In 1989, juvenile males and females grew more in both seasons (rainy and dry) than adults; for 1990, there were no differences by season or between age classes. These results are interesting since in the 1989 and 1990 rainy seasons, practically the same orders of prey and the greatest abundance of prey available in the environment were registered. A possible explanation could be that predation was more intense in 1990 than in 1989. There is little evidence that food, temperature, and humidity affect growth rates of A. nebulosus, refuting our predictions. This is mainly due to the low variation in growth observed in 1990. Therefore we think that the growth of this species reflects a complex combination of ecological and genetic factors.

Effects of traditional agroecosystems and grazing areas on amphibian diversity in a region of central Mexico.

Habitat loss or degradation due to land cover change is regarded as one of the main drivers of amphibian decline; therefore, it is imperative to assess the effects of land-cover change on this group of vertebrates. In this study, we analyze changes in alpha and beta diversity of amphibian communities found in five land-cover types: mountain cloud forest, tropical evergreen forest, shade coffee, milpa huasteca, and grazing areas; six samples sites were established for each land-cover type, separated at least one km away. The study was conducted in the northwest part of the state of Hidalgo, in a transition zone between the Sierra Madre Oriental and the Gulf of Mexico, which is a region rich in amphibian species. The results indicate that alpha diversity decreases with loss of canopy cover, this being high in mountain cloud forest, tropical evergreen forest, and Shade coffee, and low in milpa huasteca and grazing areas. The land-cover type with the highest species evenness was found in milpa huasteca and the lowest in. The highest beta diversity was observed among tropical evergreen forest and grazing areas. Mountain cloud forest contains both exclusive species and the highest number of species currently regarded as threatened by national and international conservation assessment systems. In order to preserve amphibian diversity in the study area it is vital to protect the last remnants of native vegetation, especially mountain cloud forest, but also including Shade coffee, since the latter habitat harbors amphibian diversity similar to that found in native forests. Finally, implementation of policies that both reduce Grazing areas and increase their productivity is also necessary, since these highly modified areas turn out to be the ones that affect amphibian diversity the most.

A new species of forest snake of the genus Rhadinaea from Tropical Montane Rainforest in the Sierra Madre del Sur of Oaxaca, Mexico (Squamata, Dipsadidae).

Content of the dipsadid genus Rhadinaea has changed considerably since Myers' 1974 revision. Three species groups are recognized currently in the genus. Our fieldwork in Oaxaca in June 2018 produced a single specimen of Rhadinaea considered to represent a new taxon. This new species is described from converted Premontane Wet Forest in the municipality of Santa Catarina Juquila in the Sierra Madre del Sur of southern Oaxaca, Mexico. It is most closely related to Rhadinaealaureata, from which it can be distinguished easily by color pattern and scutellation, and represents a species group distinct from the other three occupying the genus.

ResúmenLa conformación del género Rhadinaea ha cambiado considerablemente desde la revisión de Myers en 1974. Tres grupos de especies son reconocidos actualmente en el género. Nuestro trabajo de campo en Oaxaca en junio de 2018 produjo un espécimen de Rhadinaea que consideramos que representa un nuevo taxón. Describimos esta nueva especie de bosque húmedo premontano en el Municipio de Santa Catarina Juquila en la Sierra Madre del Sur en el sur de Oaxaca, México. Esta especie está más estrechamente relacionada con Rhadinaealaureata, de la cual se puede distinguir fácilmente por medio del patrón de color y escutelación, y representa un grupo distinto a los tres que forman parte del género.

Amphibian life history in a temperate environment of the Mexican Plateau: dimorphism, phenology and trophic ecology of a hylid frog, Hyla eximia (=Dryophytes eximius).

The study of demographic and life history aspects of an organism provides valuable information for its conservation. Here, we analyze the phenology of the Mountain Treefrog Hyla eximia (= Dryophytes eximius) in a temperate environment of the Mexican Plateau. Females were larger in snout-vent length and body mass than males. The peak period of activity occurred in the rainy season (May-September), with amplexus and egg deposition occurring between June and July, and larval development from July to August. A logistic model best explained observed male growth patterns, while the Von Bertalanffy model better described female growth. Notably, males grew faster than females, although females reached a larger overall body size. The diet of this species is made up of 10 prey categories. The index of diet importance indicated that males feed mainly on Coleoptera and Diptera, while females feed on Coleoptera, Diptera, Hemiptera, and Aranea. Both females and males showed a significant abundance of plant material in their stomachs, suggesting that H. eximia might exhibit highly specialized feeding behavior. Reproduction was seasonal, and both female and male reproductive cycles are synchronized with the rainy season. These natural history characteristics provide information to better understand their responses to environmental conditions.

Habitat use in eight populations of Sceloporus grammicus (Squamata: Phrynosomatidae) from the Mexican Plateau.

Studies on habitat use have often helped explain observed variation in morphology, behavior and reproductive characteristics among populations within a single species. Here we analyze morphological and ecological characteristics of individuals from the Sceloporus grammicus species complex from 7 different localities (CER, El Cerezo; PAC, Pachuca; HUI, Huichapan; EZA, Emiliano Zapata; SMR, San Miguel Regla; LMJ, La Mojonera; and LMZ, La Manzana) in the state of Hidalgo, and one locality (Cahuacán) in the State of Mexico. A canonical correspondence analysis (CCA) showed that females from PAC, EZA, LMZ, HUI, SMR and CAH populations use similar microhabitats characterized mostly by bare soil, in females from LMJ and CER use microhabitats characterized primarily by vegetation and rocks. Females were observed using 12 different types of perches. With regard to perch height use, the CCA showed that females from PAC, LMJ, LMZ, SMR, CER and CAH populations were correlated with height to nearest perch (HNP), in the rest of the females were not related to any perch use variable. In contrast, the CCA showed that males from PAC, LMJ and CAH were characterized by microhabitats with higher vegetal coverage, while males from LMZ and CER used microhabitats composed of bare soil, but males from HUI and SMR populations used microhabitats composed chiefly of bare soil and rocks. With respect to perch height use, the CCA showed that males from PAC, LMJ, EZA and LMZ were correlated with distance to the nearest perch, but the rest of the males were not correlated with any perch use variables. Males were observed in 9 different perch types. The males were larger than the females in all morphological variables analyzed. Moreover, in both sexes the snout-vent length is positively correlated with all morphological variables, and although both the slope and ordinate of the origin of all morphological variables were larger in males than females, the analysis of covariance indicated that there is no increase in the morphological variables with increasing SVL between sexes. Our results suggest that variation in habitat use and morphology among populations is an adaptive response (phenotypic plasticity) to the environmental conditions where these populations of Sceloporus grammicus occur.

Beta Diversity in a Highly Heterogeneous Area: Disentangling Species and Taxonomic Dissimilarity for Terrestrial Vertebrates.

Quantifying differences in species composition among communities provides important information related to the distribution, conservation and management of biodiversity, especially when two components are recognized: dissimilarity due to turnover, and dissimilarity due to richness differences. The ecoregions in central Mexico, within the Mexican Transition Zone, have outstanding environmental heterogeneity and harbor huge biological richness, besides differences in the origin of the biota. Therefore, biodiversity studies in this area require the use of complementary measures to achieve appropriate information that may help in the design of conservation strategies. In this work we analyze the dissimilarity of terrestrial vertebrates, and the components of turnover and richness differences, among six ecoregions in the state of Hidalgo, central Mexico. We follow two approaches: one based on species level dissimilarity, and the second on taxonomic dissimilarity. We used databases from the project "Biodiversity in the state of Hidalgo". Our results indicate that species dissimilarity is higher than taxonomic dissimilarity, and that turnover contributes more than richness differences, both for species and taxonomic total dissimilarity. Moreover, total dissimilarity, turnover dissimilarity and the dissimilarity due to richness differences were positively related in the four vertebrate groups. Reptiles had the highest values of dissimilarity, followed by mammals, amphibians and birds. For reptiles, birds, and mammals, species turnover was the most important component, while richness differences had a higher contribution for amphibians. The highest values of dissimilarity occurred between environmentally contrasting ecoregions (i.e., tropical and temperate forests), which suggests that environmental heterogeneity and differences in the origin of biotas are key factors driving beta diversity of terrestrial vertebrates among ecoregions in this complex area.

Reproductive cycles and reproductive strategies among populations of the Rose-bellied Lizard Sceloporus variabilis (Squamata: Phrynosomatidae) from central Mexico.

Species with wide distribution, generally show variations in life history characteristics, which can be attributed to environmental causes. In this study, we analyzed the reproductive cycle and reproductive characteristics from three populations (Atlapexco, San Pablo Tetlapayac, and Santa Catarina) of the lizard Sceloporus variabilis from central Mexico. The specific goal of this study was to evaluate life history characteristics such as reproductive period extent, SVL (snout-vent length) at sexual maturity, clutch size, egg mass and volume, and RCM (relative clutch mass). The San Pablo Tetlapayac population showed a larger clutch size, RCM, egg mass, and a smaller SVL, body mass and reproductive period (January-September), as well as egg volume than the Atlapexco and Santa Catarina populations. Reproductive cycle and reproductive characteristics were more similar between the Atlapexco and Santa Catarina populations. Differences found in the population of San Pablo Tetlapayac with respect to the Atlapexco and Santa Catarina populations could be attributed to environmental variations where lizard populations occur. Differences in the reproductive period and reproductive characteristics in each population could be the result of both historical (phylogenetic; e.g., reproductive mode) and nonhistorical (environmental; e.g., temperature, food availability) causes. This study showed that populations of the same species are under different selection pressures, and these affect the reproductive characteristics of populations. Our results also indicate that long-term and targeted studies on predation, use and selection of food, are needed to determine the causes of these variations in populations of S. variabilis.

Glassfrog embryos hatch early after parental desertion.

Both parental care and hatching plasticity can improve embryo survival. Research has found that parents can alter hatching time owing to a direct effect of care on embryogenesis or via forms of care that cue the hatching process. Because parental care alters conditions critical for offspring development, hatching plasticity could allow embryos to exploit variation in parental behaviour. However, this interaction of parental care and hatching plasticity remains largely unexplored. We tested the hypothesis that embryos hatch early to cope with paternal abandonment in the glassfrog Hyalinobatrachium fleischmanni (Centrolenidae). We conducted male-removal experiments in a wild population, and examined embryos' response to conditions with and without fathers. Embryos hatched early when abandoned, but extended development in the egg stage when fathers continued care. Paternal care had no effect on developmental rate. Rather, hatching plasticity was due to embryos actively hatching at different developmental stages, probably in response to deteriorating conditions without fathers. Our experimental results are supported by a significant correlation between the natural timing of abandonment and hatching in an unmanipulated population. This study demonstrates that embryos can respond to conditions resulting from parental abandonment, and provides insights into how variation in care can affect selection on egg-stage adaptations.

Oogenesis and ovarian histology in two populations of the viviparous lizard Sceloporus grammicus (Squamata: Phrynosomatidae) from the central Mexican Plateau.

The annual histological changes in ovarian morphology (oogenesis, follicular atresia, and corpus luteum) are described for the Mexican lizard Sceloporus grammicus, in two populations that inhabit contrasting environments (vegetation categories, climate, precipitation, and temperature) from Hidalgo State, Mexico. Two germinal beds were situated on the dorsal surface of each ovary of this species. In both the populations, oogenesis involves two major processes: previtellogenesis and vitellogenesis. The histological changes during previtellogenesis are similar to those for other reptilian sauropsids, whereas vitellogenesis differs and the features of this last process are described for the first time. In early previtellogenesis, primary oocytes have fibrillar chromosomes and the ooplasm stains slightly. The primordial follicles are surrounded by a granulosa composed of cuboidal follicular cells. During late previtellogenesis, the oocyte had an eccentric nucleus with lamp-brush chromosomes and multiple nucleoli. The granulosa becomes multilayered and polymorphic, containing three cell types: small, intermediate, and pyriform. The zona pellucida was homogeneous and clearly observed. In early vitellogenesis, the oocyte showed several small acidophilic granules distributed in the center and the periphery of the oocyte. As vitellogenesis progresses, the yolk platelets move toward the central area of the oocyte and they fuse to form acidophilic and homogeneous yolk. Lipid droplets were distributed irregularly in the ooplasm of the oocyte. In Zacualtipán, the results revealed a strong seasonal reproductive activity. Females had vitellogenic follicles from July to September, and pregnant females were founded from September to March. In Tizayuca, the results showed an unusual pattern of reproductive activity. Females with vitellogenic follicles and pregnant females were found throughout the year, indicating continuous reproduction. We suggest that the observed differences in reproductive activity from these populations indicate adaptative fine tuning in response to local environmental conditions. These results contribute to the knowledge of variation in vitellogenesis and reproductive strategies of this species and among spiny lizards overall.

Atypical reproductive cycles in a population of Sceloporus grammicus (Squamata: Phrynosomatidae) from the Mexican Plateau.

The spiny lizard Sceloporus grammicus (Squamata: Phrynosomatidae) is a small reptile from central México and the southern United States, occurring in a wide geographic area characterized by extensive variation in topographic and climatic regimes. Genetic variation among lineages from central México is substantial, though the extent to which this variation corresponds with life-history traits remains obscure. To address part of this puzzle, we studied a population of S. grammicus from Tepeapulco, Hidalgo, México. Male-biased sexual dimorphism was extensive in this population; males were larger than females overall, and expressed proportionately larger heads and longer limbs. Minimum size at sexual maturity was similar in the sexes (males: 43 mm; females: 42 mm). In contrast to other populations from the Central Plateau, reproductive activity of males and females was synchronous. Testicular recrudescence of adult males was initiated in October-November, and maximum testis size maintained from December to July. Female reproductive activity showed no clear seasonal pattern: females had vitellogenic follicles from October to July, and pregnant females were found throughout the year. Female body size was not related to litter size. Neither male nor female gonadal mass was correlated with any abiotic environmental variable examined. Differences in reproductive characteristics among populations of S. grammicus might be indicative of plasticity in response to local environmental conditions, local adaptation, or complex gene × environment interactions. We consider these results in the context of previously studied populations of S. grammicus from the Central Plateau and elsewhere, and propose directions for future research.