Comparative multi-locus assessment of modern Asian newts ( Cynops, Paramesotriton, and Pachytriton: Salamandridae) in southern China suggests a shared biogeographic history.

Evolutionary biologists are always interested in deciphering the geographic context of diversification, therefore they introduced the concept of comparative phylogeography, which helps to identify common mechanisms that contribute to shared genetic structures among organisms from the same region. Here, we used multi-locus genetic data along with environmental data to investigate shared phylogeographic patterns among three Asian-endemic newt genera, Cynops, Paramesotriton and Pachytriton, which occurred in montane/submontane streams or ponds in southern China. Our 222 samples from 78 localities covered the entire range of the three genera and represented the largest dataset of this group to date. We reconstructed matrilineal genealogies from two protein-coding, mitochondrial genes, and gene network from two nuclear genes. We also estimated divergence times of major cladogenetic events and used occurrence data to evaluate niche difference and similarity between lineages. Our results revealed a common basal split in all three genera that corresponds to the separation of two geographic terrains of southern China. Those ancient divergence occurred during middle to late Miocene and likely correlate with paleoclimatic fluctuations caused by the uplift of the Qinghai-Xizang (Tibet) Plateau (QTP). Particularly, the strengthening and weakening of Asian summer monsoons during the Miocene may have profoundly impacted southern China and led to repeatedly vicariance in those newts. However, despite differences in realized niches between lineages, there is no evidence for divergence of fundamental niches. Preservation of old newt matriline lineages in mountains of southern China suggests that the region acts as both museums and cradles of speciation. Based on those results, we advocate a multi-pronged protection strategy for newts in the three genera.

Tracking, Synthesizing, and Sharing Global Batrachochytrium Data at AmphibianDisease.org.

Emerging infectious diseases have been especially devastating to amphibians, the most endangered class of vertebrates. For amphibians, the greatest disease threat is chytridiomycosis, caused by one of two chytridiomycete fungal pathogens Batrachochytrium dendrobatidis (Bd) and Batrachochytrium salamandrivorans (Bsal). Research over the last two decades has shown that susceptibility to this disease varies greatly with respect to a suite of host and pathogen factors such as phylogeny, geography (including abiotic factors), host community composition, and historical exposure to pathogens; yet, despite a growing body of research, a comprehensive understanding of global chytridiomycosis incidence remains elusive. In a large collaborative effort, Bd-Maps was launched in 2007 to increase multidisciplinary investigations and understanding using compiled global Bd occurrence data (Bsal was not discovered until 2013). As its database functions aged and became unsustainable, we sought to address critical needs utilizing new technologies to meet the challenges of aggregating data to facilitate research on both Bd and Bsal. Here, we introduce an advanced central online repository to archive, aggregate, and share Bd and Bsal data collected from around the world. The Amphibian Disease Portal (https://amphibiandisease.org) addresses several critical community needs while also helping to build basic biological knowledge of chytridiomycosis. This portal could be useful for other amphibian diseases and could also be replicated for uses with other wildlife diseases. We show how the Amphibian Disease Portal provides: (1) a new repository for the legacy Bd-Maps data; (2) a repository for sample-level data to archive datasets and host published data with permanent DOIs; (3) a flexible framework to adapt to advances in field, laboratory, and informatics technologies; and (4) a global aggregation of Bd and Bsal infection data to enable and accelerate research and conservation. The new framework for this project is built using biodiversity informatics best practices and metadata standards to ensure scientific reproducibility and linkages across other biological and biodiversity repositories.

Forever young: Linking regeneration and genome size in salamanders.

BACKGROUND: Salamanders stand out among vertebrate animals in two key characteristics: their ability to regenerate body parts, and their large and variable genome sizes. RESULTS: Here we show how to unite seemingly disparate facets of salamander biology, regeneration ability, and genome size variation, into one synthetic view. Large and variable genome sizes may be the key to understanding the prodigious ability of most salamanders to regenerate damaged or lost body parts. We report a correlate of genome size variation that has been previously neglected: the impacts of genome size on the structure and function of the genes themselves. Salamanders are, in essence, paradoxically much younger, especially at the cellular level than their chronological age would suggest. CONCLUSIONS: Because of the large size and range of variation in genome size in salamanders, we hypothesize that this relationship uncouples a dynamic interaction between growth and differentiation in processes of morphogenesis, pattern formation, and regeneration in ways that are unique among vertebrates.

Two extremely rare new species of fossorial salamanders of the genus Oedipina (Plethodontidae) from northwestern Ecuador.

We describe two new species of salamanders of the genus Oedipina, subgenus Oedopinola, from two localities on the northwestern foothills of Ecuador, at elevations between 921 and 1,067 m. These are the southernmost members of the genus. We examined different museum collections and we found just three specimens of Oedipina from Ecuador, obtained throughout the history of herpetological collections in the country. We identify two of the three specimens as new species, but refrain from assigning a specific identity to the third, pending further study. Oedipina villamizariorum sp. n. is a medium-sized member of the genus, with a narrow, relatively pointed head and blunt snout; dorsolaterally oriented eyes, moderate in size; and digits that are moderately long and having pointed tips. Oedipina ecuatoriana sp. n., somewhat larger, has a narrow head and broadly rounded snout; this new species differs from all known Oedipina by the distinctive presence of paired prefrontal bones and a reduced phalangeal formula: 0-0-1-0; 0-1-2-1-1. We provide detailed descriptions of the osteology of both new species. Finally, we present a phylogenetic hypothesis for the genus, including one of the two new species, based on partial sequences of mitochondrial DNA.

Taxonomic status of the Neotropical salamanders Bolitoglossa altamazonica and Bolitoglossa peruviana (Amphibia: Caudata: Plethodontidae), with the description of a new species from Northern Peru.

We examine the phylogenetic relationships among salamanders of the genus Bolitoglossa (Eladinea) distributed in the Amazonian basin of northern Peru and southern Ecuador and assess species diversity based on morphological and phylogenetic analyses. We infer a molecular phylogeny using sequences from two mitochondrial (Cytb, 16S) and two nuclear genes (RAG-1, POMC). We find two well-supported subclades, one including [B. altamazonica + B. peruviana] + B. awajun sp. n., and the other including Bolitoglossa sp. Ituxi + Bolitoglossa sp. Jurúa. Ecuadorian lineages form divergent clades from the Peruvian lineages. Accordingly, Ecuadorian populations previously assigned to Bolitoglossa peruviana sensu lato are treated as members of a Bolitoglossa equatoriana species complex. A newly defined Bolitoglossa altamazonica species complex contains only populations from the Amazonian rainforest of Peru. Maximum likelihood and Bayesian Inference analyses confirm the phylogenetic placement of B. altamazonica and B. peruviana, and support recognition of a related new species of Bolitoglossa. The uncorrected genetic distances between the new species and B. altamazonica are 6.5% for Cytb and 4.9% for 16S; and the uncorrected genetic distances between the new species and B. peruviana are 8.0% for Cytb and 3.9% for 16S. Additionally, analyses of nuclear gene sequences show no haplotype sharing between the new species and closely related species. The new species is distinguished from its congeners by a combination of the following morphological characters: (1) Standard length mean 37.7 mm in males (range 32.0-42.2; n=5) and 41.4 mm in females (range 34.9-48.2; n=6); (2) in life, dorsal coloration uniformly brown with a dark brown triangular marking between the eyes or some irregular light cream spots or patches on the head, back and flanks; (3) iris pale golden; (4) in preservative, dark brown venter with cream mottlings or moderate-sized blotches on the gular region, belly, cloacal region and tail; (5) tips of third finger and third toe protuberant and pointed with nearly complete webbing on the hands and feet; (6) 11-26 maxillary teeth and 8-24 vomerine teeth. Given that the syntypes of B. altamazonica are lost, we designate a neotype for B. altamazonica from Allpahuayo Mishana National Reserve, Loreto department, Peru. Newly collected specimens from ~30 km NE from Moyobamba (type locality of B. peruviana) provide a better understanding of B. peruviana and enable us to show that it is the sister taxon of B. altamazonica. The new species is known from pre-montane forests in Cordillera Escalera Regional Conservation Area, Cordillera Azul National Park and Shucshuyacu, San Martin department, Peru at 485-1311 m elevation,  ~75 km SE from Moyobamba. Bolitoglossa awajun sp. n. is the fourth endemic species of salamander from Peru.

Slender salamanders (genus Batrachoseps) reveal Southern California to be a center for the diversification, persistence, and introduction of salamander lineages.

BACKGROUND: The southern California biodiversity hotspot has had a complex geological history, with both plate tectonic forces and sea level changes repeatedly reconfiguring the region, and likely driving both lineage splittings and extinctions. Here we investigate patterns of genetic divergence in two species of slender salamanders (Plethodontidae: Batrachoseps) in this region. The complex geological history in combination with several organismal traits led us to predict that these species harbor multiple ancient mitochondrial lineages endemic to southern California. These species belong to a clade characterized by fine-scale mitochondrial structure, which has been shown to track ancient splits. Both focal species, Batrachoseps major and B. nigriventris, are relatively widely distributed in southern California, and estimated to have persisted there across millions of years. Recently several extralimital populations of Batrachoseps were found in the San Joaquin Valley of California, a former desert area that has been extensively modified for agriculture. The origins of these populations are unknown, but based on morphology, they are hypothesized to result from human-mediated introductions of B. major. METHODS: We sequenced the mitochondrial gene cytochrome b from a geographically comprehensive sampling of the mitochondrial lineages of B. major and B. nigriventris that are endemic to southern California. We used phylogenetic analyses to characterize phylogeographic structure and identify mitochondrial contact zones. We also included the San Joaquin Valley samples to test whether they resulted from introductions. We used a bootstrap resampling approach to compare the strength of isolation-by-distance in both Batrachoseps species and four other salamander species with which they co-occur in southern California. RESULTS: The northern lineage of B. major harbors at least eight deeply differentiated, geographically cohesive mitochondrial subclades. We identify geographic contact between many of these mtDNA lineages and some biogeographic features that are concordant with lineage boundaries. Batrachoseps nigriventris also has multiple deeply differentiated clades within the region. Comparative analyses highlight the smaller spatial scales over which mitochondrial divergence accumulates in Batrachoseps relative to most other salamander species in southern California. The extralimital populations of Batrachoseps from the San Joaquin Valley are assigned to B. major and are shown to result from at least two independent introductions from different source populations. We also suggest that B. major on Catalina Island, where it is considered native, may be the result of an introduction. Some of the same traits that facilitate the build-up of deep phylogeographic structure in Batrachoseps likely also contribute to its propensity for introductions, and we anticipate that additional introduced populations will be discovered.

Descriptions of five new species of the salamander genus Chiropterotriton (Caudata: Plethodontidae) from eastern Mexico and the status of three currently recognized taxa.

The genus Chiropterotriton is endemic to Mexico with a geographical distribution along the Sierra Madre Oriental, the Trans Mexican Volcanic Belt and the Sierra de Juárez. The recent use of molecular tools has shown that Mexico's amphibian diversity is highly underestimated, including a large number of cryptic, unnamed species. Chiropterotriton has 18 described species including terrestrial, arboreal and cave-dwelling species. In previous molecular studies, the presence of multiple undescribed species was evident. We present a phylogenetic hypothesis based on mitochondrial data, which includes all described species and six undescribed taxa. Based on the morphological analyses and, when available, combined with molecular data, we describe five new species of the genus; Chiropterotriton casasi sp. nov., C. ceronorum sp. nov., C. melipona sp. nov., C. perotensis sp. nov. and C. totonacus sp. nov. In addition, we redescribe two others: Chiropterotriton chiropterus and C. orculus, and provide a comparable account of one additional sympatric congener. This increases the number of species in the genus to 23, which represent a considerable component of Mexican plethodontid richness.

Taxonomic revision of black salamanders of the Aneides flavipunctatus complex (Caudata: Plethodontidae).

We present a taxonomic revision of the black salamander (Aneides flavipunctatus) complex of northwestern California and extreme southeastern Oregon. The revision is based on a number of published works as well as new molecular and morphological data presented herein. The subspecies Aneides flavipunctatus niger Myers & Maslin 1948 is raised in rank to a full species. It is isolated far to the south of the main range on the San Francisco Peninsula, south and west of San Francisco Bay. Another geographically isolated set of populations occurs well inland in Shasta County, northern CA, mainly in the vicinity of Shasta Lake. It is raised from synonymy and recognized as Aneides iecanus (Cope 1883). The remaining taxa occur mainly along and inland from the coast from the vicinity of the Russian River and Lake Berryessa/Putah Creek, north to the vicinity of the Smith River near the Oregon border and more inland along the Klamath and Trinity Rivers and tributaries into Oregon. The northern segment of this nearly continuous range is named Aneides klamathensis Reilly and Wake 2019. We use molecular data to provide a detailed examination of a narrow contact zone between the northern A. klamathensis and the more southern A. flavipunctatus in southern Humboldt County in the vicinity of the Van Duzen and main fork of the Eel rivers. To the south is the remnant of the former species and it takes the name Aneides flavipunctatus (Strauch 1870). It is highly diversified morphologically and genetically and requires additional study.

Amphibians.

Since the dawn of history, amphibians have been a part of human culture. Western Europeans built fires for cooking and warmth, adding large logs as needed. What occasionally emerged was astounding: large black animals (which had found shelter in the logs) with four legs and a tail, jet black with striking bright yellow spots. These fire salamanders were variously thought to be the product of the fire itself, or, as Aristotle reported, capable of extinguishing fire. Pliny the Elder is said to have tested this idea by throwing a salamander into flames - the salamander died! - nevertheless the association with fire persisted. Pliny perpetuated other fantastical claims, which spread; even Leonardo da Vinci contributed to the legend, and myths from different regions merged - at one point, asbestos was claimed to be salamander wool. Salamanders were attributed great powers; a single salamander upstream was thought to be sufficient to kill an army. King Francis I. of France chose a salamander as his emblem - a powerful symbol, born of fire, filled with poison, immune from burning, and even able to douse flames. Before the emergence of great cities and conurbations, people grew up surrounded by nature. Salamanders and newts, toads and frogs were all part of normal human experience. Myths such as those surrounding the fire salamanders were commonplace. Shakespeare's witches brewed with an eye of newt and tail of frog. As a child, we raised tadpoles and were taught to shudder at the appearance of a tiger salamander in a root cellar. In general, amphibians are seen as benign and harmless, even helpful as creatures that devour harmful insects and serve as an alternative food source. Thus, it came as a shock to most biologists and to the public at large in the 1980s that amphibians around the world were in decline and that they were at greater risk of extinction as a taxon than any other vertebrate group. A study of every amphibian species known in 2004 showed that on the order of 40% were at high risk of extinction, and by 2008, the decline of amphibians was seen as evidence of an impending sixth mass extinction.

Selection and environmental adaptation along a path to speciation in the Tibetan frog Nanorana parkeri.

Tibetan frogs, Nanorana parkeri, are differentiated genetically but not morphologically along geographical and elevational gradients in a challenging environment, presenting a unique opportunity to investigate processes leading to speciation. Analyses of whole genomes of 63 frogs reveal population structuring and historical demography, characterized by highly restricted gene flow in a narrow geographic zone lying between matrilines West (W) and East (E). A population found only along a single tributary of the Yalu Zangbu River has the mitogenome only of E, whereas nuclear genes of W comprise 89-95% of the nuclear genome. Selection accounts for 579 broadly scattered, highly divergent regions (HDRs) of the genome, which involve 365 genes. These genes fall into 51 gene ontology (GO) functional classes, 14 of which are likely to be important in driving reproductive isolation. GO enrichment analyses of E reveal many overrepresented functional categories associated with adaptation to high elevations, including blood circulation, response to hypoxia, and UV radiation. Four genes, including DNAJC8 in the brain, TNNC1 and ADORA1 in the heart, and LAMB3 in the lung, differ in levels of expression between low- and high-elevation populations. High-altitude adaptation plays an important role in maintaining and driving continuing divergence and reproductive isolation. Use of total genomes enabled recognition of selection and adaptation in and between populations, as well as documentation of evolution along a stepped cline toward speciation.

A new terrestrial species of Chiropterotriton (Caudata: Plethodontidae) from central Mexico.

Chiropterotriton is a relatively small genus that comprises 15 species with great morphological and ecological diversity. In previous studies, molecular data provided evidence for a considerable number of species that remain undescribed. In this study, we describe one new species, Chiropterotriton chico sp. nov. based on molecular and morphological characters. We present mtDNA phylogenetic analyses using Bayesian inference and maximum likelihood that include all described and several undescribed species. Morphometric data from eight recognized species provide evidence for the distinctiveness of the new taxon. Description of this new species adds to the already high salamander diversity of the state of Hidalgo, which is an important area for the diversification of the genus.

Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary.

Frogs (Anura) are one of the most diverse groups of vertebrates and comprise nearly 90% of living amphibian species. Their worldwide distribution and diverse biology make them well-suited for assessing fundamental questions in evolution, ecology, and conservation. However, despite their scientific importance, the evolutionary history and tempo of frog diversification remain poorly understood. By using a molecular dataset of unprecedented size, including 88-kb characters from 95 nuclear genes of 156 frog species, in conjunction with 20 fossil-based calibrations, our analyses result in the most strongly supported phylogeny of all major frog lineages and provide a timescale of frog evolution that suggests much younger divergence times than suggested by earlier studies. Unexpectedly, our divergence-time analyses show that three species-rich clades (Hyloidea, Microhylidae, and Natatanura), which together comprise ∼88% of extant anuran species, simultaneously underwent rapid diversification at the Cretaceous-Paleogene (K-Pg) boundary (KPB). Moreover, anuran families and subfamilies containing arboreal species originated near or after the KPB. These results suggest that the K-Pg mass extinction may have triggered explosive radiations of frogs by creating new ecological opportunities. This phylogeny also reveals relationships such as Microhylidae being sister to all other ranoid frogs and African continental lineages of Natatanura forming a clade that is sister to a clade of Eurasian, Indian, Melanesian, and Malagasy lineages. Biogeographical analyses suggest that the ancestral area of modern frogs was Africa, and their current distribution is largely associated with the breakup of Pangaea and subsequent Gondwanan fragmentation.

Biology of tiny animals: three new species of minute salamanders (Plethodontidae: Thorius) from Oaxaca, Mexico.

We describe three new species of minute salamanders, genus Thorius, from the Sierra Madre del Sur of Oaxaca, Mexico. Until now only a single species, T. minutissimus, has been reported from this region, although molecular data have long shown extensive genetic differentiation among geographically disjunct populations. Adult Thorius pinicola sp. nov., T. longicaudus sp. nov., and T. tlaxiacus sp. nov. are larger than T. minutissimus and possess elliptical rather than oval nostrils; T. pinicola and T. longicaudus also have longer tails. All three new species occur west of the range of T. minutissimus, which has the easternmost distribution of any member of the genus. The new species are distinguished from each other and from other named Thorius in Oaxaca by a combination of adult body size, external morphology and osteology, and by protein characters (allozymes) and differences in DNA sequences. In addition, we redescribe T. minutissimus and a related species, T. narisovalis, to further clarify the taxonomic status of Oaxacan populations and to facilitate future studies of the remaining genetically differentiated Thorius that cannot be satisfactorily assigned to any named species. Populations of all five species considered here appear to have declined dramatically over the last one or two decades and live specimens are difficult to find in nature. Thorius may be the most endangered genus of amphibians in the world. All species may go extinct before the end of this century.

Enlarged Multilocus Data set Provides Surprisingly Younger Time of Origin for the Plethodontidae, the Largest Family of Salamanders.

Deep phylogenetic relationships of the largest salamander family Plethodontidae have been difficult to resolve, probably reflecting a rapid diversification early in their evolutionary history. Here, data from 50 independent nuclear markers (total 48,582 bp) are used to reconstruct the phylogeny and divergence times for plethodontid salamanders, using both concatenation and coalescence-based species tree analyses. Our results robustly resolve the position of the enigmatic eastern North American four-toed salamander (Hemidactylium) as the sister taxon of Batrachoseps + Tribe Bolitoglossini, thus settling a long-standing question. Furthermore, we statistically reject sister taxon status of Karsenia and Hydromantes, the only plethodontids to occur outside the Americas, leading us to new biogeographic hypotheses. Contrary to previous long-standing arguments that plethodontid salamanders are an old lineage originating in the Cretaceous (more than 90 Ma), our analyses lead to the hypothesis that these salamanders are much younger, arising close to the K-T boundary (~66 Ma). These time estimates are highly stable using alternative calibration schemes and dating methods. Our data simulation highlights the potential risk of making strong arguments about phylogenetic timing based on inferences from a handful of nuclear genes, a common practice. Based on the newly obtained timetree and ancestral area reconstruction results, we argue that (i) the classic "Out of Appalachia" hypothesis of plethodontid origins is problematic; (ii) the common ancestor of extant plethodontids may have originated in northwestern North America in the early Paleocene; (iii) origins of Eurasian plethodontids likely result from two separate dispersal events from western North America via Beringia in the late Eocene (~42 Ma) and the early Miocene (~23 Ma), respectively.

An integrative approach to phylogeography: investigating the effects of ancient seaways, climate, and historical geology on multi-locus phylogeographic boundaries of the Arboreal Salamander (Aneides lugubris).

BACKGROUND: Phylogeography is an important tool that can be used to reveal cryptic biodiversity and to better understand the processes that promote lineage diversification. We studied the phylogeographic history of the Arboreal Salamander (Aneides lugubris), a wide-ranging species endemic to the California floristic province. We used multi-locus data to reconstruct the evolutionary history of A. lugubris and to discover the geographic location of major genetic breaks within the species. We also used species distribution modeling and comparative phylogeography to better understand the environmental factors that have shaped the genetic history of A. lugubris. RESULTS: We found six major mitochondrial clades in A. lugubris. Nuclear loci supported the existence of at least three genetically distinct groups, corresponding to populations north of the San Francisco Bay and in the Sierra Nevada, in the Santa Cruz Mountains, and in the central coast and southern California. All of the genetic breaks in mitochondrial and nuclear loci corresponded to regions where historical barriers to dispersal have been observed in other species. Geologic or water barriers likely were the most important factors restricting gene flow among clades. Climatic unsuitability during glacial maximum may have contributed to the isolation of the mitochondrial clades in the central coast and southern California. A projection of our species distribution model to a future scenario with a moderate amount of climate change suggests that most of the range of A. lugubris will remain climatically suitable, but climatic conditions in the Sierra Nevada and low elevation areas in Southern California are likely to deteriorate. CONCLUSIONS: Aneides lugubris contains substantial cryptic genetic diversity as a result of historical isolation of populations. At least two (and perhaps three) evolutionarily significant units in A. lugubris merit protection; all six mitochondrial clades should be considered as management units within the species.

Ring species as demonstrations of the continuum of species formation.

In the mid-20th century, Ernst Mayr (1942) and Theodosius Dobzhansky (1958) championed the significance of 'circular overlaps' or 'ring species' as the perfect demonstration of the gradual nature of species formation. As an ancestral species expands its range, wrapping around a geographic barrier, derived taxa within the ring display interactions typical of populations, such as genetic and morphological intergradation, while overlapping taxa at the terminus of the ring behave largely as sympatric, reproductively isolated species. Are ring species extremely rare or are they just difficult to detect? What conditions favour their formation? Modelling studies have attempted to address these knowledge gaps by estimating the biological parameters that result in stable ring species (Martins et al. 2013), and determining the necessary topographic parameters of the barriers encircled (Monahan et al. 2012). However, any generalization is undermined by a major limitation: only a handful of ring species are known to exist in nature. In addition, many of them have been broken into multiple species presumed to be evolving independently, usually obscuring the evolutionary dynamics that generate diversity. A paper in this issue of Molecular Ecology by Fuchs et al. (2015), focused on the entire genealogy of a bulbul (Alophoixus) species complex, offers key insights into the evolutionary processes underlying diversification of this Indo-Malayan bird. Their findings fulfil most of the criteria that can be expected for ring species (Fig. ): an ancestor has colonized the mainland from Sundaland, expanded along the forested habitat wrapping around Thailand's lowlands, adjacent taxa intergrade around the ring distribution, and terminal taxa overlap at the ring closure. Although it remains unclear whether ring divergence has resulted in restrictive gene flow relative to that observed around the ring, their results suggest that circular overlaps might be more common in nature than currently recognized in the literature. Most importantly, this work shows that the continuum of species formation that Mayr and Dobzhansky praised in circular overlaps is found in biological systems currently described as 'rings of species', in addition to the idealized 'ring species'.

Indeterminate Growth: Could It Represent the Ancestral Condition?

Although we are used to the idea that many organisms stop growing when they reach a predictable size, in many taxa, growth occurs throughout the life of an organism, a phenomenon referred to as indeterminate growth. Our comparative analysis suggests that indeterminate growth may indeed represent the ancestral condition, whereas the permanent arrest of growth may be a more derived state. Consistent with this idea, in diverse taxa, the basal branches show indeterminate growth, whereas more derived branches arrest their growth. Importantly, in some closely related taxa, the termination of growth has evolved in mechanistically distinct ways. Also, even within a single organism, different organs can differ with respect to whether they terminate their growth or not. Finally, the study of tooth development indicates that, even at the level of a single tissue, multiple determinate patterns of growth can evolve from an ancestral one that is indeterminate.