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Article Is Available Online Pod Morphology Before and After the Permian Mass Extinction At Biogeosciences, 15, 5489–5502, 2018 https://doi.org/10.5194/bg-15-5489-2018 © Author(s) 2018. This work is distributed under the Creative Commons Attribution 4.0 License. Significance of climate and hydrochemistry on shape variation – a case study on Neotropical cytheroidean Ostracoda Claudia Wrozyna1,2, Thomas A. Neubauer3,4, Juliane Meyer1, Maria Ines F. Ramos5, and Werner E. Piller1 1Institute of Earth Sciences, NAWI Graz Geocenter, University of Graz, 8010 Graz, Austria 2Institute for Geophysics and Geology, University of Leipzig, 04109 Leipzig, Germany 3Department of Animal Ecology & Systematics, Justus Liebig University, 35392 Giessen, Germany 4Naturalis Biodiversity Center, Leiden, 2300 RA, the Netherlands 5Coordenação de Ciências da Terra e Ecologia, Museu Paraense Emílio Goeldi, 66077-830, Brazil Correspondence: Claudia Wrozyna ([email protected]) Received: 13 September 2017 – Discussion started: 10 November 2017 Revised: 21 August 2018 – Accepted: 24 August 2018 – Published: 14 September 2018 Abstract. How environmental change affects a species’ phe- ality, annual precipitation and chloride and sulfate concen- notype is crucial not only for taxonomy and biodiversity as- trations. We suggest that increased temperature seasonality sessments but also for its application as a palaeo-ecological slowed down growth rates during colder months, potentially and ecological indicator. Previous investigations addressing triggering the development of shortened valves with well- the impact of the climate and hydrochemical regime on os- developed brood pouches. Differences in chloride and sul- tracod valve morphology have yielded contrasting results. fate concentrations, related to fluctuations in precipitation, Frequently identified ecological factors influencing carapace are considered to affect valve development via controlling shape are salinity, cation, sulfate concentrations, and alkalin- osmoregulation and carapace calcification. The factors iden- ity. Here, we present a thorough approach integrating data tified by our analyses represent hitherto unknown drivers for with the carapace outline and surface details of the ubiqui- ostracod ecophenotypy in other species and therefore suggest tous Neotropical cytheroidean ostracod species Cytheridella that environmental predictors for morphological variability ilosvayi, as well as several climatic and hydrochemical vari- are not consistent across non-marine ostracods. ables, in order to investigate a potential link between mor- phology and environmental conditions. A recent study pre- viously demonstrated considerable biogeographical variation in valve morphology among Floridian, Mexican and Brazil- 1 Introduction ian populations of this species. We hypothesize that the cli- matic differences between the regions it inhabits and as- Understanding how species respond to environmental change sociated differences in hydrochemical regimes have influ- is crucial for their application as proxies for past climate enced valve morphology and eventually led to biogeographi- fluctuations as well as in forecasting the future dynamics cally distinctive groups. Generalized least-squares Procrustes and distributions of species. Morphological diversity repre- analyses based on outline and structural features were ap- sents a key feature of the interpretation of faunal changes plied to the left and right valves of adult females and males. (Wagner and Erwin, 2006) and ecological shifts (Mahler et The analyses identified relative carapace length and shape al., 2010) and prompts discussions about speciation and ex- symmetry as most important morphological characteristics tinction processes over time (e.g. Ciampaglio, 2004). Differ- representing shape differences across all datasets. Two-block ences in shape and size among species have been shown to partial least-squares analyses and multiple regressions indi- relate to changes of environmental parameter, in particular, cate strong relationships between morphological and envi- differences in temperature across various clades (e.g. Loehr ronmental variables, specifically with temperature season- et al., 2010; Maan and Seehausen, 2011; Danner and Green- berg, 2015). Within freshwater invertebrates, ecophenotypic Published by Copernicus Publications on behalf of the European Geosciences Union. 5490 C. Wrozyna et al.: Significance of climate and hydrochemistry on shape variation response has been documented for a variety of species, both the biogeographic variability in C. ilosvayi are well under- recent and fossil (e.g. Hellberg et al., 2001; Zieritz and stood, the causes for the regional differences have not been Aldridge, 2009; Inoue et al., 2013; Neubauer et al., 2013; investigated. We hypothesize that the climatic differences be- Clewing et al., 2015). tween the regions inhabited by Cytheridella ilosvayi and the Ostracods represent a popular proxy group for climate and associated differences in hydrochemical regimes have influ- ecosystem changes due to their occurrence in various habi- enced valve morphology and finally led to biogeographically tats, ranging from most inland waters to marine and inter- distinctive groups. We apply two-block partial least-squares stitial and even semiterrestrial and terrestrial environments analyses and multiple regression analyses in order to test for (e.g. Horne, 2005). Their distribution is controlled by eco- covariation between the two sets of parameters (morphol- logical factors such as salinity, temperature, and ion com- ogy, environment) and to identify the morphological char- position of the ambient water (e.g. Ruiz et al., 2013). The acteristics and environmental variables that contribute most study of ecophenotypic variation in response to environmen- to the relationship. This approach is new to the research of tal change (Anadón et al., 2002; Frenzel et al., 2012; Fürsten- ecophenotypy in ostracods for the following reasons: the in- berg et al., 2015; van der Meeren et al., 2010) demonstrates vestigation of the relationships between shape variations and another approach using ostracods for palaeoenvironmental environmental conditions is based on (1) a dataset covering studies. Due to their calcitic valves, they have an excellent a supra-regional geographical scale that coincides with the fossil record and are utilized as palaeoenvironmental and geographical range of Cytheridella, (2) environmental vari- biostratigraphic indicators (Anadón et al., 2002). A number ables including local hydrochemical data and regional cli- of studies have shown that ornamentation, noding, sieve pore matic data, (3) morphometric data that are based on a combi- shape, and carapace size are linked to environmental factors, nation of landmarks and semilandmarks, including the cara- e.g. salinity, temperature, water depth, and nutrient availabil- pace outline and positions of pore tubules on the surface ity (van Harten, 1975; Yin et al., 1999; Majoran et al., 2000; of the carapaces, and (4) the identification of environmental van Harten, 2000; Anadón et al., 2002; Frenzel and Boomer, variables that covary with morphological changes are based 2005; Medley et al., 2007; Marco-Barba et al., 2013; Meyer on multivariate statistical analyses. et al., 2016; Boomer et al., 2017). Especially with the rise of morphometric techniques, investigations dealing with cara- pace shape variation in relation to environmental variables 2 Material and methods have also increased (Yin et al., 1999; Baltanas et al., 2002; 2.1 Material van der Meeren et al., 2010; Ramos et al., 2017; Grossi et al., 2017). Yet, the use of morphological data, even those Specimens of C. ilosvayi derive from several sampling cam- based on morphometric analyses (Baltanas et al., 2002, 2003; paigns in Florida, Mexico, and Brazil during 2009–2015 van der Meeren et al., 2010; Grossi et al., 2017), has been (Fig. 1). A detailed list of the sampled localities is avail- restricted to either landmark-based or outline-based studies able in Table S1. Only adult valves were utilized in this but has rarely used a combination of both (e.g. Ramos et al., study, providing a sufficient number of left and right valves 2017). Few studies integrate geographic gradients into their for both sexes. Right and left valves were investigated sep- statistical analyses and corresponding climate variables or arately due to dimorphism in size and shape (Wrozyna et a reduced number of predictor variables. Moreover, shape– al., 2014). Aside from this, females and males were anal- environment relationships are commonly identified based on ysed separately because a large part of inter-valve variation simple linear regressions or qualitative observations on mul- has been shown to depend on sexual differences (such as the tivariate ordination methods. presence of brood pouches in females; Wrozyna et al., 2016). Here, we apply a thorough approach integrating morpho- metric data from carapace outline and surface details, as well 2.2 Predictor variables as several climatic and hydrochemical variables, in order to investigate a potential link between morphology and environ- Altogether, 15 variables were included in the analyses. Si- mental conditions. The subject of study is the valves of the multaneously to water sampling, field variables (electrical Neotropical cytheroidean ostracod species, Cytheridella ilos- conductivity, water temperature and pH) were measured in vayi. Studies by von Daday (1905) and Wrozyna et al. (2016, situ at all sample sites using a WTW multi-sensor probe 2018a) already demonstrated considerable
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