1 Introduction

Molecular systematics has emerged as a pivotal tool in understanding the evolutionary relationships and biodiversity of invertebrates. By analyzing genetic material, researchers can uncover the phylogenetic relationships among species, which often remain obscured by morphological similarities or differences. This approach has been particularly transformative in the study of symbiotic relationships, such as those between coral reef invertebrates and their endosymbiotic dinoflagellates, revealing complex evolutionary histories and adaptive radiations. Additionally, molecular systematics has provided insights into the evolutionary and plastic responses of freshwater invertebrates to climate change, highlighting the role of genetic and phenotypic plasticity in adaptation (Stoks et al., 2013). The integration of molecular data with fossil records has also been crucial in reconciling discrepancies between molecular and morphological evolutionary rates, offering a more comprehensive understanding of macroevolutionary patterns (Sansom and Wills, 2013; Condamine et al., 2016).

Geological changes, such as climatic shifts and tectonic events, have profoundly influenced the evolution and distribution of invertebrates. For instance, the Miocene-Pliocene transition, characterized by significant climatic changes and low CO₂ levels, triggered adaptive radiations in symbiotic dinoflagellates, which in turn affected their invertebrate hosts. Similarly, the reduction in glacier cover due to global warming has led to consistent changes in the functional diversity and community assembly of river invertebrates across different biogeographic regions (Brown et al., 2017). These geological changes not only impact the physical environment but also drive evolutionary processes, leading to diversification and adaptation in invertebrate populations. The fossil record, despite its biases, provides critical data for understanding these macroevolutionary transitions and the impact of past environmental changes on invertebrate diversification (Sansom and Wills, 2013; Condamine et al., 2016).

This study aims to analyze the phylogenetic relationships among invertebrate species using molecular data; examine the impact of historical geological events, such as climatic shifts and tectonic activities, on the diversification and adaptation of invertebrates; integrate molecular and fossil data to provide a comprehensive understanding of invertebrate evolution; and identify key functional traits that mediate the responses of invertebrates to environmental changes and their effects on ecosystem services. This study hope to enhance the understanding of the evolutionary dynamics of invertebrates and their resilience to geological changes, thereby contributing to the broader field of evolutionary biology and conservation.

2 Geological Changes and Their Impact on Invertebrate Evolution

2.1 Major geological events (e.g., plate tectonics, volcanic activity)

Geological events such as plate tectonics and volcanic activity have played a significant role in shaping the evolutionary pathways of invertebrates. The movement of tectonic plates can lead to the formation of new habitats and the isolation of populations, which in turn drives speciation and diversification. For instance, the Cenomanian-Turonian succession in Wadi Tarfa, Egypt, reveals how sea-level fluctuations influenced the community structure of benthic invertebrates, with different species thriving during transgressive and regressive phases (Abdelhady et al., 2020). Volcanic activity can also create new landforms and alter existing habitats, providing new ecological niches for invertebrates to exploit.

2.2 Climate shifts and habitat transformations

Climate shifts have had profound impacts on invertebrate evolution by altering habitats and the availability of resources. During the last deglacial warming, for example, there was a significant shift in the mollusk fauna from cold-tolerant species to warmth-adapted species in the Chinese Loess Plateau. This shift was closely linked to changes in vegetation, highlighting the interdependency between plant and invertebrate communities (Dong et al., 2020). Similarly, freshwater invertebrates have shown both evolutionary and plastic responses to climate change, with changes in phenology and body size being driven by temperature increases (Stoks et al., 2013). These examples underscore the importance of climate as a driver of evolutionary change in invertebrates.

2.3 Fossil evidence and evolutionary milestones

Fossil records provide crucial insights into the evolutionary milestones of invertebrates in response to geological changes. The fossil evidence from the Cenomanian-Turonian period in Wadi Tarfa illustrates how invertebrate communities adapted to long-term sea-level changes, with different species dominating during various phases of transgression and regression (Abdelhady et al., 2020). Additionally, the study of ice-binding proteins (IBPs) in intertidal invertebrates suggests that these proteins evolved multiple times in response to freezing conditions, highlighting the role of environmental pressures in driving molecular evolution (Box et al., 2022). These findings demonstrate how fossil evidence can help researchers understand the evolutionary responses of invertebrates to past geological and climatic events.

3 Advances in Molecular Systematics

3.1 Techniques in DNA sequencing and genomics

The advent of next-generation sequencing (NGS) has revolutionized the field of molecular systematics by significantly reducing the cost and time required for sequencing, thereby enabling the generation of large-scale genomic data. This has facilitated the inclusion of multiple genes and even whole genomes in phylogenetic studies, providing a more comprehensive understanding of evolutionary relationships. For instance, the use of shotgun sequencing to assemble mitochondrial genomes from complex ecological mixtures has proven effective in overcoming taxonomic impediments and expanding the phylogenetic representation of various lineages (Crampton-Platt et al., 2015). Additionally, the development of exon-capture systems has allowed for the generation of extensive data matrices from museum samples, further enhancing the robustness and comprehensiveness of phylogenetic trees (O’hara et al., 2017). These advancements underscore the transformative impact of NGS on molecular systematics, enabling more detailed and accurate reconstructions of evolutionary histories.

3.2 Phylogenetic analysis and evolutionary trees

Phylogenetic analysis has greatly benefited from the integration of molecular data, leading to more robust and statistically supported evolutionary trees. The use of multilocus analyses and genome-scale data has challenged traditional views and provided new insights into the relationships among various taxa (Young and Gillung, 2020). For example, the phylogenetic reconstruction of New Zealand cheilostome bryozoans using sequences from 17 genes revealed discrepancies between molecular and morphological data, highlighting the need for rethinking current systematic hypotheses (Orr et al., 2021). Moreover, the application of phylogenetic comparative methods, supported by null-hypothesis testing and power analysis via simulation, has increased the confidence and robustness of these analyses (Pyron, 2015). These methodological advancements have significantly enhanced our ability to infer evolutionary relationships and understand the diversification of life.

3.3 Integration of molecular data with morphological and paleontological data

The integration of molecular data with morphological and paleontological evidence provides a more holistic view of evolutionary history. This total evidence approach allows for the resolution of deep branching patterns and the empirical testing of tree reconstruction techniques. By combining molecular sequences with morphological observations, researchers can discriminate between conflicting hypotheses and evaluate the assumptions underlying phylogenetic methods. For instance, the integration of molecular and morphological data in the study of cheilostome bryozoans has provided insights into the evolutionary history of specific traits, such as the presence of frontal shields (Orr et al., 2021). Additionally, paleontological data offer independent means of calibrating molecular trees, thereby providing insights into rates of molecular evolution in the geological past. This comprehensive approach enhances the accuracy and reliability of phylogenetic reconstructions, contributing to a more complete understanding of the Tree of Life.

The advances in DNA sequencing and genomics, coupled with sophisticated phylogenetic analysis techniques and the integration of diverse data types, have significantly advanced the field of molecular systematics. These developments have enabled more detailed and accurate reconstructions of evolutionary histories, providing deeper insights into the diversification and relationships of invertebrates in response to geological changes.

4 Invertebrate Case Studies in Geological Contexts

4.1 Arthropods: evolution and diversification

Arthropods represent the most diverse animal phylum, with their evolutionary history and phylogenetic relationships being subjects of extensive research. Molecular phylogenetics has established arthropods as monophyletic, placing them within the ecdysozoans, a clade of molting animals that includes nematodes and other phyla. This molecular framework has clarified relationships within major arthropod groups, such as the Pancrustacea, which includes insects and crustaceans. The evolutionary history of arthropods is further illuminated by a rich fossil record, which, despite some conflicts in analyses, provides crucial insights into their origins and diversification. Molecular time-trees, calibrated with fossils, estimate the origins of arthropods in the Ediacaran period, with most deep nodes dating to the Cambrian. Early stem-group arthropods, such as lobopodians, were worm-like animals with annulated appendages, highlighting the significant morphological evolution within the phylum (Figure 1) (Giribet and Edgecombe, 2019).

4.2 Mollusks: phylogenetic relationships and adaptations

Mollusks, including snails, octopuses, and clams, exhibit a remarkable diversity of body plans, second only to arthropods in species number. Despite extensive study, their evolutionary relationships remain poorly resolved, with significant questions about the origins and morphological evolution within the group. Recent phylogenomic studies have generated transcriptome data for various molluscan species, supporting the clade Aculifera, which includes groups with spicules but no true shells, and the monophyly of Conchifera. These studies have also clarified relationships among major molluscan groups, such as the sister group relationship between Scaphopoda (tusk shells) and Gastropoda. This well-resolved phylogenetic tree provides a framework for further studies on mollusc evolution, development, and anatomy (Smith et al., 2011). Additionally, the interaction between mollusks and their parasites, such as Schistosoma mansoni, reveals complex mechanisms of immune diversification and adaptation, further illustrating the evolutionary dynamics within this phylum (Roger et al., 2008).

4.3 Echinoderms: molecular insights into evolutionary responses

Echinoderms, as one of the most primitive deuterostomes, offer valuable insights into the evolutionary biology of chordates due to their close phylogenetic relationship. Studies on the N-glycomic capacity of echinoderms, such as the brittle star Ophiactis savignyi, have revealed a diverse set of glycoconjugates, including structures reminiscent of both vertebrates and invertebrates. This diversity in glycosylation positions echinoderms at an evolutionary nexus between invertebrates and vertebrates, highlighting their unique molecular adaptations (Figure 2) (Eckmair et al., 2020). Furthermore, the crystallographic data on echinoderm skeletons have been used as taxonomic tools, providing insights into their evolutionary history and classification. These molecular and structural studies underscore the adaptive responses of echinoderms to their environments and their evolutionary significance.

5 Molecular Markers and Genetic Diversity

5.1 Types of molecular markers used in systematics

Molecular markers are essential tools in systematics for assessing genetic diversity and understanding evolutionary relationships among species. Various types of molecular markers have been employed, each with its specific applications and advantages. Commonly used markers include:

Microsatellites (SSRs): These are short, repetitive DNA sequences that are highly polymorphic and widely used for population genetic studies due to their high mutation rates and codominant inheritance (Kindie, 2021; Yi et al., 2023).

Single nucleotide polymorphisms (SNPs): SNPs are single base-pair variations in the DNA sequence. They are abundant throughout the genome and provide high-resolution data for genetic diversity and population structure analysis (Steele and Pires, 2011; Yi et al., 2023; Wenne, 2023).

Amplified fragment length polymorphisms (AFLPs): AFLPs are used for DNA fingerprinting and assessing genetic variation. They are particularly useful for species with limited genomic information (Kindie, 2021).

Random amplified polymorphic DNA (RAPD): RAPD markers are used for genetic mapping and diversity studies. They are quick and cost-effective but have lower reproducibility compared to other markers (Kindie, 2021).

Mitochondrial DNA (mtDNA): mtDNA markers, such as cytochrome b and control region sequences, are commonly used for phylogenetic studies and species identification due to their maternal inheritance and relatively rapid mutation rates (Arif and Khan, 2009).

5.2 Assessing genetic variation and population structure

Assessing genetic variation and population structure is crucial for understanding the evolutionary processes and ecological dynamics of species. Molecular markers provide insights into the genetic diversity within and between populations, which is essential for conservation and management strategies.

Genetic diversity analysis: Molecular markers like SSRs and SNPs are used to quantify genetic diversity, which reflects the adaptive potential of populations. High genetic diversity is often associated with greater resilience to environmental changes (Steele and Pires, 2011; Kindie, 2021; Yi et al., 2023).

Population structure: Markers such as AFLPs and mtDNA are used to infer population structure and gene flow. This information helps in identifying distinct population units and understanding their evolutionary history (Arif and Khan, 2009; Yi et al., 2023).

Phylogeography: Molecular markers enable the study of historical population movements and demographic changes. This is particularly important for species affected by past climatic events and habitat fragmentation (Steele and Pires, 2011; Li et al., 2020; Yi et al., 2023).

5.3 Implications for conservation and biodiversity

The application of molecular markers in conservation biology has significant implications for the preservation of biodiversity. By providing detailed genetic information, these markers help in formulating effective conservation strategies.

Conservation units: Molecular markers aid in defining conservation units by identifying genetically distinct populations and cryptic species. This ensures that conservation efforts are directed towards preserving genetic diversity (Li et al., 2020; Yi et al., 2023).

Management strategies: Genetic data from molecular markers inform management practices such as habitat restoration, captive breeding, and reintroduction programs. This helps in maintaining genetic integrity and adaptive potential of populations (Li et al., 2020; Kindie, 2021; Yi et al., 2023).

Monitoring and assessment: Continuous monitoring of genetic diversity using molecular markers allows for the assessment of conservation interventions' effectiveness. This dynamic approach helps in adapting strategies to changing environmental conditions and threats (Li et al., 2020; Yi et al., 2023).

In conclusion, molecular markers are indispensable tools in the field of systematics and conservation biology. They provide critical insights into genetic diversity, population structure, and evolutionary history, which are essential for the effective conservation and management of biodiversity.

6 Speciation and Phylogeography

6.1 Mechanisms of speciation in invertebrates

Speciation in invertebrates is driven by a variety of mechanisms, both genetic and ecological. Genetic mechanisms include repeated founder events, hybridization, and sexual selection, which have been observed in Hawaiian terrestrial arthropods. Ecological mechanisms, such as shifts in habitat and host affiliation, also play a significant role in the diversification of these species. In the case of freshwater copepod crustaceans in European lakes, deep mitochondrial splits among populations suggest that divergence of lineages predates the Pleistocene glaciations, indicating that historical and biogeographical factors significantly shape modern patterns of distribution (Kochanova et al., 2021). Additionally, the intense uplifting of the Qinghai-Tibetan Plateau and Quaternary climate oscillations have driven speciation and genetic structure in the Odorrana graminea sensu lato in Southern China (Chen et al., 2019).

6.2 Phylogeographic patterns in relation to geological changes

Geological changes have a profound impact on the phylogeographic patterns of invertebrates. In the Hawaiian archipelago, repeated colonization of new island groups has led to lineages progressing down the island chain, with the most ancestral groups on the oldest islands. Similarly, in New Zealand, phylogeographic studies reveal signatures of partitioning in various regions and expansion in different directions, influenced by Pliocene tectonic events and range expansion following the last glacial maximum (Trewick et al., 2011). In southeastern Australia, the interaction between physiogeographic landscape context and life history characteristics, particularly dispersal ability, has generated predictable outcomes for how species responded to Pleistocene climatic changes (Garrick et al., 2012). In unglaciated eastern North America, recurrent phylogeographic patterns, such as the Appalachian Mountain discontinuity and the Mississippi River discontinuity, are attributable to isolation and differentiation during Pleistocene glaciation.

6.3 Case studies: examples of speciation events

Several case studies illustrate speciation events in invertebrates. In the Hawaiian terrestrial arthropods, speciation patterns are classified into three categories: single representatives of a lineage throughout the islands, species radiations with single endemic species on different volcanoes or islands, and single widespread species within a radiation of species that exhibit local endemism. In Tallaganda, southeastern Australia, flightless low-mobility forest invertebrates, such as springtails and terrestrial flatworms, exhibit spatial patterns of intraspecific genetic diversity that conform to topography-based divisions, highlighting cases of phylogeographic congruence and incongruence (Garrick et al., 2012). In European freshwater copepod crustaceans, deep mitochondrial splits among populations indicate that divergence of lineages predates the Pleistocene glaciations, suggesting that historical and biogeographical factors significantly shape modern patterns of distribution (Kochanova et al., 2021). Lastly, in Southern China, the Odorrana graminea sensu lato exhibits five major highly divergent lineages, with phylogenetic analyses revealing significant gene flow events and demographic expansions during the last glacial maximum (Chen et al., 2019).

7 Adaptation and Survival Strategies

7.1 Genetic adaptations to environmental stressors

Invertebrates exhibit a range of genetic adaptations that enable them to survive and thrive in response to various environmental stressors. For instance, freshwater invertebrates have shown genetic changes in response to climate change, particularly in traits related to temperature and photoperiod adjustments (Stoks et al., 2013). Similarly, studies on pollution adaptation have highlighted the complex genetic responses to heavy metal contamination, although the evidence for consistent adaptive responses across different species remains mixed (Loria et al., 2019). Additionally, DNA methylation has been identified as a mechanism by which invertebrates can adjust to novel environments, such as urban settings, by altering stress response genes (Holdt et al., 2022). These genetic adaptations are crucial for maintaining homeostasis and ensuring survival in changing environments.

7.2 Evolution of reproductive and developmental strategies

The evolution of reproductive and developmental strategies in invertebrates is a key aspect of their adaptation to environmental changes. Phenological shifts, such as changes in the timing of life cycle events, have been observed in response to climate change, driven by both genetic and plastic responses (Schilthuizen and Kellermann, 2013). In some cases, maladaptive plasticity can occur when ancestral developmental systems are co-opted to meet new environmental challenges, leading to reduced fitness. However, genetic compensation mechanisms can remodel these plastic responses to restore fitness (Velotta and Cheviron, 2018). This remodeling process is often one of the earliest steps in the adaptive evolution of reproductive and developmental strategies, allowing invertebrates to better cope with novel stressors.

7.3 Phenotypic plasticity and behavioral adaptations

Phenotypic plasticity plays a significant role in the survival of invertebrates under environmental stress. This plasticity allows organisms to express different phenotypes depending on environmental conditions, thereby increasing their fitness and expanding their ecological niches (Hoffmann and Bridle, 2021). For example, thermal plasticity in insects enables them to adjust their developmental and physiological traits in response to temperature fluctuations, which is particularly relevant in the context of climate change (Rodrigues and Beldade, 2020). Behavioral adaptations, such as optimal foraging and thermoregulation, also contribute to the ability of invertebrates to cope with environmental variability (Hoffmann and Bridle, 2021). However, the effectiveness of these plastic responses can vary, and in some cases, they may be maladaptive, necessitating further evolutionary adjustments (Ho and Zhang, 2018).

In summary, the adaptation and survival strategies of invertebrates in response to geological changes involve a complex interplay of genetic adaptations, evolution of reproductive and developmental strategies, and phenotypic plasticity. These mechanisms collectively enable invertebrates to navigate and thrive in dynamic and often challenging environments.

8 Case Studies in Molecular Systematics

8.1 Case study 1: invertebrate response to volcanic activity

Volcanic activity can have profound impacts on invertebrate populations, primarily through changes in habitat and food availability. For instance, the study of terrestrial mollusks in the Chinese Loess Plateau during the last deglacial warming period provides insights into how invertebrates respond to significant environmental changes. The research demonstrated a shift from cold-tolerant to warmth-adapted mollusk species, indicating that volcanic activity and subsequent climatic changes can drive significant shifts in invertebrate community composition (Figure 3) (Dong et al., 2020). This case study highlights the importance of understanding the indirect impacts of geological events on invertebrate populations through changes in vegetation and habitat structure.

8.2 Case study 2: marine invertebrates and sea level changes

Marine invertebrates are highly sensitive to sea level fluctuations, which can alter their habitats and community structures. A study on the Cenomanian-Turonian succession in Wadi Tarfa, Egypt, revealed that sea-level changes significantly influenced the community structure of benthic invertebrates. The research identified three distinct invertebrate communities associated with different stages of sea-level changes: transgression, maximum flooding, and highstand system tracts. These communities exhibited varying diversity and dominance patterns, with higher diversity during maximum flooding due to increased environmental stability and substrate heterogeneity (Abdelhady et al., 2020). This case study underscores the critical role of sea-level changes in shaping marine invertebrate communities over geological timescales.

8.3 Case study 3: impact of glaciation on terrestrial invertebrates

Glaciation events have historically impacted terrestrial invertebrate populations by altering their habitats and food sources. The study of brachiopod communities during the Late Paleozoic Ice Age in Bolivia provides valuable insights into how glaciation affects invertebrate diversity and community structure. The research found that genus richness was higher during glacial periods, likely due to smaller body sizes and time-averaged mixing of genera from different depths. Additionally, the study observed a monotonic increase in warm-water genera and North American biogeographic affinity, suggesting that community changes were driven by the northward drift of Bolivia rather than glacial cycles (Badyrka et al., 2013). This case study highlights the complex interactions between glaciation, habitat changes, and invertebrate community dynamics.

9 Future Directions and Challenges

9.1 Emerging technologies in molecular systematics

The field of molecular systematics is rapidly evolving with the advent of new technologies. One promising area is the use of multi-omics approaches, which integrate data from genomics, transcriptomics, and metabolomics to provide a comprehensive understanding of species' responses to environmental changes. For instance, a study on Antarctic marine invertebrates used metabolomics and transcriptomics to uncover species-specific molecular responses to acute warming, highlighting the complexity and diversity of these responses (Clark et al., 2016). Additionally, advancements in conservation genomics are crucial for preserving the biodiversity of marine invertebrates, which represent a significant portion of animal biodiversity. These technologies can help identify novel genes and genomic innovations that are essential for adaptation and survival (Lopez et al., 2019).

9.2 Integrative approaches to studying invertebrate evolution

Integrative approaches that combine phylogenetic systematics with other methodologies are essential for a deeper understanding of invertebrate evolution. Phylogenetic systematics has already provided new insights into the life history evolution of marine invertebrates, challenging traditional assumptions and revealing the diversity of reproductive and developmental modes. Moreover, the use of DNA barcoding and integrative species delimitation has proven effective in identifying cryptic species and understanding the evolutionary relationships among invertebrates. For example, a study on photosynthetic sea slugs used DNA barcoding and morphological data to identify multiple candidate species within what were previously considered single species, demonstrating the utility of integrative approaches in uncovering hidden biodiversity (Krug et al., 2013).

9.3 Conservation and management implications

The findings from molecular systematics and integrative approaches have significant implications for conservation and management. Understanding the molecular and genetic basis of species' responses to environmental stressors, such as microplastic ingestion, can inform conservation strategies. A systematic review of the antioxidant system in invertebrates exposed to microplastics highlighted the need for more comprehensive studies to identify the specific characteristics of microplastics that cause oxidative stress (Trestrail et al., 2020). Additionally, the study of benthic invertebrate communities in response to sea-level fluctuations provides valuable insights into how long-term environmental changes affect biodiversity and ecosystem stability, which is crucial for conservation planning (Abdelhady et al., 2020). Finally, the development of best practices in conservation genomics, including genomic monitoring and profiling, can enhance broader conservation goals by providing baseline data for in situ and ex situ conservation efforts (Lopez et al., 2019).

10 Concluding Remarks

The study of molecular systematics in invertebrates has revealed significant insights into how these organisms have responded to geological changes over time. Molecular systematics has shown that endosymbiotic dinoflagellates, particularly Symbiodinium spp., have undergone significant adaptive radiation since the Miocene-Pliocene transition. This diversification is linked to host specialization and allopatric differentiation, suggesting a response to major climatic changes and low CO₂ levels during that period. Freshwater invertebrates exhibit both evolutionary and plastic responses to climate change. While there is evidence of phenotypic plasticity and genetic changes, the extent of these adaptations varies, with some traits showing more robust responses to temperature changes. The analysis of functional traits in invertebrates has highlighted the importance of key functional traits that mediate the effects of biodiversity on ecosystem services. These traits are crucial for understanding how invertebrates respond to environmental changes and contribute to ecosystem functioning.

The reduction in glacier cover has led to consistent increases in the functional diversity of river invertebrates across multiple biogeographic regions. This pattern is driven by dispersal limitation and environmental filtering, indicating predictable mechanisms governing community responses to environmental changes. The fossilization process can distort evolutionary trees by causing taxa to appear more primitive than they are. This bias, known as stem-ward slippage, affects the interpretation of macroevolutionary rates and sequences, highlighting the need for careful consideration of fossil data in evolutionary studies. Insect diversification patterns show both early bursts and steady rates of diversification, with major shifts occurring in holometabolous orders. This suggests that clade-specific innovations, rather than broad environmental changes, have driven diversification in insects. The molecular phylogeny of New Zealand cheilostome bryozoans has provided a framework for testing systematic hypotheses. The study found that lower taxonomic levels are robust, while higher-level systematics require rethinking. The presence of frontal shields in cheilostomes was also linked to diversification rates.

The findings from these studies have broader implications for evolutionary biology: The adaptive radiation of symbiotic dinoflagellates and freshwater invertebrates in response to climatic changes underscores the dynamic nature of evolutionary processes. These examples highlight the importance of understanding how environmental factors drive diversification and adaptation in different ecosystems. Identifying key functional traits that link biodiversity to ecosystem services is crucial for predicting how ecosystems will respond to future environmental changes. This approach can inform conservation strategies and ecosystem management practices. The recognition of biases in the fossil record, such as stem-ward slippage, is essential for accurate phylogenetic reconstruction and understanding macroevolutionary patterns. This awareness can improve the integration of fossil and molecular data in evolutionary studies. The study of insect diversification and bryozoan phylogeny highlights the role of clade-specific innovations and morphological traits in driving evolutionary processes. These findings contribute to a more nuanced understanding of the factors that influence diversification across different taxonomic groups.

Future research should focus on continuing efforts to reconcile molecular and fossil evidence will enhance researchers’ understanding of evolutionary timelines and diversification patterns. Further studies on functional traits across diverse invertebrate groups will help identify key traits that influence ecosystem services. This knowledge is vital for developing robust indicators of ecosystem health and resilience. As climate change continues to affect ecosystems globally, it is important to monitor and understand the evolutionary and plastic responses of invertebrates. Long-term studies and experimental approaches will provide deeper insights into these adaptive processes. Many invertebrate groups remain understudied in terms of their molecular systematics. Expanding research to include these taxa will provide a more comprehensive picture of invertebrate evolution and biodiversity. By addressing these future prospects, researchers can continue to unravel the complex interactions between invertebrates and their changing environments, contributing to the broader field of evolutionary biology.

Acknowledgements

The authors thank the two anonymous peer reviewers for their careful review and feedback on the manuscript of this study, which helped us to improve and refine this study.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

References 

Abdelhady A., Mohamed R., Fathy D., and Ali A., 2020, Benthic invertebrate communities as a function of sea-level fluctuations and hydrodynamics: A case from the Cenomanian-Turonian of Wadi Tarfa (Eastern Desert, Egypt), Journal of African Earth Sciences, 168: 103870.

https://doi.org/10.1016/j.jafrearsci.2020.103870

Arif I., and Khan H., 2009, Molecular markers for biodiversity analysis of wildlife animals: a brief review, Animal Biodiversity and Conservation, 32(1): 9-17.

https://doi.org/10.32800/abc.2009.32.0009

Badyrka K., Clapham M., and López S., 2013, Paleoecology of brachiopod communities during the late Paleozoic ice age in Bolivia (Copacabana Formation, Pennsylvanian-Early Permian), Palaeogeography, Palaeoclimatology, Palaeoecology, 387: 56-65.

https://doi.org/10.1016/j.palaeo.2013.07.016

Box I., Matthews B., and Marshall K., 2022, Molecular evidence of intertidal habitats selecting for repeated ice-binding protein evolution in invertebrates, Journal of Experimental Biology, 225(Suppl_1): jeb243409.

https://doi.org/10.1242/jeb.243409
PMid:35258616

Brown L., Khamis K., Wilkes M., Blaen P., Brittain J., Carrivick J., Fell S., Friberg N., Füreder L., Gíslason G., Hainie S., Hannah D., James W., Lencioni V., Ólafsson J., Robinson C., Saltveit S., Thompson C., and Milner A., 2017, Functional diversity and community assembly of river invertebrates show globally consistent responses to decreasing glacier cover, Nature Ecology & Evolution, 2: 325-333.

https://doi.org/10.1038/s41559-017-0426-x
PMid:29255301

Chen Z., Li H., Zhai X., Zhu Y., He Y., Wang Q., Li Z., Jiang J., Xiong R., and Chen X., 2019, Phylogeography, speciation and demographic history: contrasting evidence from mitochondrial and nuclear markers of the Odorrana graminea sensu lato (Anura, Ranidae) in China, Molecular Phylogenetics and Evolution, 144: 106701.

https://doi.org/10.1016/j.ympev.2019.106701
PMid:31811937

Clark M., Sommer U., Sihra J., Thorne M., Morley S., King M., Viant M., and Peck L., 2016, Biodiversity in marine invertebrate responses to acute warming revealed by a comparative multi- omics approach, Global Change Biology, 23: 318-330.

https://doi.org/10.1111/gcb.13357
PMid:27312151 PMCid:PMC6849730

Condamine F., Clapham M., and Kergoat G., 2016, Global patterns of insect diversification: towards a reconciliation of fossil and molecular evidence? Scientific Reports, 6: 19208.

https://doi.org/10.1038/srep19208
PMid:26778170 PMCid:PMC4725974

Crampton-Platt A., Timmermans M., Gimmel M., Kutty S., Cockerill T., Khen C., and Vogler A., 2015, Soup to tree: the phylogeny of beetles inferred by mitochondrial metagenomics of a Bornean rainforest sample, Molecular Biology and Evolution, 32: 2302-2316.

https://doi.org/10.1093/molbev/msv111
PMid:25957318 PMCid:PMC4540967

Dong Y., Wu N., Jiang W., Li F., and Lu H., 2020, Cascading response of flora and terrestrial mollusks to last deglacial warming, Global Ecology and Conservation, 24: e01360.

https://doi.org/10.1016/j.gecco.2020.e01360

Eckmair B., Jin C., Karlsson N., Abed-Navandi D., Wilson I., and Paschinger K., 2020, Glycosylation at an evolutionary nexus: the brittle star Ophiactis savignyi expresses both vertebrate and invertebrate N-glycomic features, The Journal of Biological Chemistry, 295: 3173-3188.

https://doi.org/10.1074/jbc.RA119.011703
PMid:32001617 PMCid:PMC7062179

Garrick R., Rowell D., and Sunnucks P., 2012, Phylogeography of saproxylic and forest floor invertebrates from Tallaganda, south-eastern Australia, Insects, 3: 270-294.

https://doi.org/10.3390/insects3010270
PMid:26467960 PMCid:PMC4553628

Giribet G., and Edgecombe G., 2019, The phylogeny and evolutionary history of arthropods, Current Biology, 29: R592-R602.

https://doi.org/10.1016/j.cub.2019.04.057
PMid:31211983

Ho W., and Zhang J., 2018, Evolutionary adaptations to new environments generally reverse plastic phenotypic changes, Nature Communications, 9: 350.

https://doi.org/10.1038/s41467-017-02724-5
PMid:29367589 PMCid:PMC5783951

Hoffmann A., and Bridle J., 2021, The dangers of irreversibility in an age of increased uncertainty: revisiting plasticity in invertebrates, Oikos, 4: e08715.

https://doi.org/10.1111/oik.08715

Holdt B., Kartzinel R., Oers K., Verhoeven K., and Ouyang J., 2022, Changes in the rearing environment cause reorganization of molecular networks associated with DNA methylation, The Journal of Animal Ecology, 92(3): 648-664.

https://doi.org/10.1111/1365-2656.13878
PMid:36567635

Kindie B., 2021, Assess molecular marker applications for genetic variety analysis in biodiversity conservation status, Molecular Biology, 10: 287.

https://doi.org/10.37421/2168-9547.21.10.287

Kochanova E., Nair A., Sukhikh N., Väinölä R., and Husby A., 2021, Patterns of cryptic diversity and phylogeography in four freshwater copepod crustaceans in European lakes, Diversity, 13(9): 448.

https://doi.org/10.3390/d13090448

Krug P., Vendetti J., Rodriguez A., Retana J., Hirano Y., and Trowbridge C., 2013, Integrative species delimitation in photosynthetic sea slugs reveals twenty candidate species in three nominal taxa studied for drug discovery, plastid symbiosis or biological control, Molecular Phylogenetics and Evolution, 69(3): 1101-1119.

https://doi.org/10.1016/j.ympev.2013.07.009
PMid:23876292 PMCid:PMC3788053

Li Y., Liu C., Rong W., Luo S., Nong S., Wang J., and Chen X., 2020, Applications of molecular markers in conserving endangered species, Biodiversity Science, 28(3): 367-375.

https://doi.org/10.17520/biods.2019414

Lopez J., Kamel B., Medina M., Collins T., and Baums I., 2019, Multiple facets of marine invertebrate conservation genomics, Annual Review of Animal Biosciences, 7: 473-497.

https://doi.org/10.1146/annurev-animal-020518-115034
PMid:30485758

Loria A., Cristescu M., and Gonzalez A., 2019, Mixed evidence for adaptation to environmental pollution, Evolutionary Applications, 12: 1259-1273.

https://doi.org/10.1111/eva.12782
PMid:31417613 PMCid:PMC6691217

O’hara T., Hugall A., Thuy B., Stöhr S., and Martynov A., 2017, Restructuring higher taxonomy using broad-scale phylogenomics: the living Ophiuroidea, Molecular Phylogenetics and Evolution, 107: 415-430.

https://doi.org/10.1016/j.ympev.2016.12.006
PMid:27940329

Orr R., Martino E., Gordon D., Ramsfjell M., Mello H., Smith A., and Liow L., 2021, A broadly resolved molecular phylogeny of New Zealand cheilostome bryozoans as a framework for hypotheses of morphological evolution, Molecular Phylogenetics and Evolution, 161: 107172.

https://doi.org/10.1016/j.ympev.2021.107172
PMid:33813020

Pyron R., 2015, Post-molecular systematics and the future of phylogenetics, Trends in Ecology & Evolution, 30(7): 384-389.

https://doi.org/10.1016/j.tree.2015.04.016
PMid:26014093

Rodrigues Y., and Beldade P., 2020, Thermal plasticity in insects’ response to climate change and to multifactorial environments, Frontiers in Ecology and Evolution, 8: 271.

https://doi.org/10.3389/fevo.2020.00271

Roger E., Grunau C., Pierce R., Hirai H., Gourbal B., Galinier R., Emans R., Cesari I., Cosseau C., and Mitta G., 2008, Controlled chaos of polymorphic mucins in a metazoan parasite (Schistosoma mansoni) interacting with its invertebrate host (Biomphalaria glabrata), PLoS Neglected Tropical Diseases, 2(11): e330.

https://doi.org/10.1371/journal.pntd.0000330
PMid:19002242 PMCid:PMC2576457

Sansom R., and Wills M., 2013, Fossilization causes organisms to appear erroneously primitive by distorting evolutionary trees, Scientific Reports, 3: 2545.

https://doi.org/10.1038/srep02545
PMid:23985991 PMCid:PMC3756334

Schilthuizen M., and Kellermann V., 2013, Contemporary climate change and terrestrial invertebrates: evolutionary versus plastic changes, Evolutionary Applications, 7: 56-67.

https://doi.org/10.1111/eva.12116
PMid:24454548 PMCid:PMC3894898

Smith S., Wilson N., Goetz F., Feehery C., Andrade S., Rouse G., Giribet G., and Dunn C., 2011, Resolving the evolutionary relationships of molluscs with phylogenomic tools, Nature, 480: 364-367.

https://doi.org/10.1038/nature10526
PMid:22031330

Steele P., and Pires J., 2011, Biodiversity assessment: state-of-the-art techniques in phylogenomics and species identification, American Journal of Botany, 98(3): 415-425.

https://doi.org/10.3732/ajb.1000296
PMid:21613135

Stoks R., Geerts A., and Meester L., 2013, Evolutionary and plastic responses of freshwater invertebrates to climate change: realized patterns and future potential, Evolutionary Applications, 7: 42-55.

https://doi.org/10.1111/eva.12108
PMid:24454547 PMCid:PMC3894897

Trestrail C., Nugegoda D., and Shimeta J., 2020, Invertebrate responses to microplastic ingestion: reviewing the role of the antioxidant system, The Science of the Total Environment, 734: 138559.

https://doi.org/10.1016/j.scitotenv.2020.138559
PMid:32470656

Trewick S., Wallis G., and Morgan-Richards M., 2011, The invertebrate life of New Zealand: a phylogeographic approach, Insects, 2: 297-325.

https://doi.org/10.3390/insects2030297
PMid:26467729 PMCid:PMC4553545

Velotta J., and Cheviron Z., 2018, Remodeling ancestral phenotypic plasticity in local adaptation: a new framework to explore the role of genetic compensation in the evolution of homeostasis, Integrative and Comparative Biology, 58(6): 1098-1110.

https://doi.org/10.1093/icb/icy117
PMid:30272147 PMCid:PMC6658864

Wenne R., 2023, Single nucleotide polymorphism markers with applications in conservation and exploitation of aquatic natural populations, Animals, 13(6): 1089.

https://doi.org/10.3390/ani13061089
PMid:36978629 PMCid:PMC10044284

Yi S., Zeng C., Li Y., and Muangmai N., 2023, Editorial: population genetics and conservation of aquatic species, Frontiers in Genetics, 13: 1052740.

https://doi.org/10.3389/fgene.2022.1052740
PMid:36685815 PMCid:PMC9845757

Young A., and Gillung J., 2020, Phylogenomics- principles, opportunities and pitfalls of big-data phylogenetics, Systematic Entomology, 45(2): 225-247.

https://doi.org/10.1111/syen.12406